PART II
ECOLOGICAL STUDIES 9 a() 1 3 _ A 7002 2970010B 183
Foreword to Part II
As stated in Chapter 1, the second part of this dissertation entails the description of ecological studies at a site in eastern
Tasmania. In these chapters, as in the entire work, the use and expression of units is as recommended in "Style manual for authors, editors and printers of Australian Government publications" (third edition, Commonwealth of Australia, 1978). Abbreviations and symbols used are as follows:
number in sample
S.D. standard deviation
Student's t 2 X chi-squared
d.f. degrees of freedom
probability
n.s. not significant at the level p=0.05
significant at the level p=0.05
** significant at the level p=0.01
*** significant at the level p=0.001
Other abbreviations and symbols are explained in the text. 184 CHAPTER FIVE
ENVIRONMENT
A description of the Anglers Creek study site:
methods, results and discussion.
Section 5.1 Location and Access
2 The study site chosen was an area of about 3000 m of low closed-forest (Specht, 1970) near Anglers Creek, on Moaners Tier,
3 km southeast of Tooms Lake and 17 km from the east coast of Tasmania
(Figure 5.1 and Plate 5.1). The Jurassic dolerite intrusions in east- ern Tasmania have given rise to a series of low plateaus, or tiers
(Davies, 1965). Moaners Tier is a smaller example which rises to
750 in near the study site, which is at an altitude of 600 m. There 0 is a northerly slope of 5 in the study area, of which no part is closer than 20 in to Anglers Creek. The creek itself is small at this point, and is reduced to a string of pools after dry summer weather.
Tasmanian Pulp and Forest Holdings, Limited, are holders of the concession to timber in the Anglers Creek area. . A road built by this company provided access to the study site, as the site was near a side- branch of the main road (road "M"). This side road, Tower Road, bounded an area set aside by T.P. & F.H. as a flora reserve, and although clear- felling took place nearby (Plate 5.1), the study site, which was in the reserve, was not threatened.
Section 5.2 Climate
The climate at the study site could be classified on a global basis, as temperate maritime (Bureau of Meteorology, 1979), while by
Tasmanian standards, the area has mild temperatures and fairly low rain- fall. Mean temperatures in January and June are 13.8°C and 4.4 °C res- pectively at Lake Leake Chalet, 25 km to the north (Commonwealth Bureau Figure 5.1 Section of Little Swanport
1:100 000 sheet (no. 8413, edition 2)
showing Tooms Lake area and Anglers
Creek study site (black star). Solid
black line indicates route of logging
road "M" and side branch. Location
of Tooms Lake shown in Figure 4.1.
U s•- t 446. , • S , 1 s 144- 2 / ot' - -,-' ' I I/ ---
04. •
Pigeon wood ---NN Hill
or Do • 459 Goatrotk Hill
MerediiN.
• 476 K, Hi
Daylight B.re. Hill Sup/ al _ S tla flaestel Mit
MOUNT •/ • 495 TOOMS Buxtons / Lookout
va. LAKE Goannn Hill 4 464• cress CY'l° 1, Wilsons .6! - loel SAN CTUA Barren
-t
.1f
Sassafras Hill
296
Des,/ Hills
She Oak Bonk PONT ypoo, Ptrs '5vfalso'617-- SWAN/Don •mu.)an41. RIE/ER Littlo Swontiort
Black Jacks islands f;:#g)-. Hill
r, WAN:. rON
Creek
c Plate 5.1 Aerial photograph of Anglers Creek area,
November 1979 (scale 1:8000). Rectangle
indicates location of study site.
• • Plate 5.2 Aerial photograph of Anglers Creek area
during 1967. Rectangle indicates
location of study site.
ilipartment Photographic Section 134 Macquarie Street 185 of Meteorology records).
The mean annual precipitation at Tooms Lake is 665 mm, falling in an average 158 raindays spread fairly evenly through the year. The driest and wettest months are January (41 mm) and July (73 mm) res- pectively. Figure 5.2 shows the average distribution of precipitation
(Tooms Lake) and temperature (Lake Leake Chalet) throughout the year.
Precipitation at Toms Lake during the period of study (1976-1977) is shown in Figure 5.3. Snow falls occasionally in winter but rarely settles, while frosts are quite frequent during the winter months.
During the study period, certain parameters of microclimate were measured at the Anglers Creek area.
Section 5.3 Temperature Measurement
These data were collected in three different ways. A Grant
Instruments Model D, 9-channel automatic temperature recorder was used
to compare the diurnal temperature cycles in different microhabitats.
The temperature at the soil-litter interface, where most amphipods are
found, was monitored by maximum-minimum thermometer and by a sucrose inversion method.
(a) Diurnal cycle
Four thermistor probes were set up at Anglers Creek, at 20 am above ground in a shaded position, on the litter surface, under the
litter, and at a depth of 10 cm in the soil. The recorder was run
for one week in March 1976, and Figure 5.4 shows the plotted results
for 14 March, a warm autumn day following a warm period. The small diurnal variation of litter and soil temperatures is clearly shown, while
the litter surface, as would be expected, is more strongly influenced by the air temperature. It should be noted that the total diurnal fluc- tuation in air in the forest at Anglers Creek shown here is less than that recorded at the weather station at Lake Leake, where the maximum o o was 23.1 C and .the minimum 6.3 C. It is most probable that this effect qoii 'AboT alooq ow SL 6T -L S6T ) (r) 0 0 0-3 rt- IT PJ UT QJ aN p aq oNuq Z 'S alrIBTa 0 ▪ Average rainfall CD 0 w looms Lake(mm) ▪
Ul
▪ ----I CO Average temperature Ul CD Lake Leake( U1 ° C ) 1975 1976 48 - 46 - 44 -
42 - 40 -
38 -
36 - 34 -
32 - 30 -
28 -
26 -
TO 16 -
8 - 6 - 4
1,I I I h I ( II iI II I II I L i ii L 1... I DEC NOV DEC JAN FEB MAR APRIL MAY JUNE JULY AUG SEPT OC T NOV 38 1977
36
34
32
30
28
26
24 22
20
71 18
16
14
12 10
8
6
2
I ii 1J. I ,. ii .. .1. , ii I. 11. [I .1 • • P.. I hi. JAN FEB MAR APRIL MAY JUNE JULY AUG SEPT OCT NOV DEC
Figure 5.3 (both pages) Daily rainfall recorded at Tooms Lake during study period, from Bureau of Meteorology records.
■IN
• • II --;77:11 um, ■ o air, shaded
9 - • litter surface
8- o litter
7- ▪ soil, 1 cm
6-
I II I I I I I I I I II 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 Time of day (h)
Figure 5.4 Hourly temperature measurements in selected positions at Anglers Creek study site on 14 March 1976. 186 is due mainly to the insulating effect of forest canopy (Geiger, 1961).
(b) Maximum-minimum temperature measurements
A Zeal maximum-minimum thermometer was sunk slightly into the soil in the study area leaving the bulb at soil surface level, and the litter was replaced over it. This was read monthly or more frequently and the results are shown in Figure 5.5.
Cc) Sucrose inversion method
This method of temperature integration was first developed by
Pallmann et al. (1940) and further refined by Berthet (1960), while Lee
(1969) has outlined some of its limitations. Basically, an integrated temperature value for a period of time is obtained by measuring the extent to which a sucrose solution has hydrolysed, when exposed for a known number of days in the field. The speed of this reaction in a solution of a certain concentration is dependent on pH and temperature..
The pH is known because the prepared solutions are buffered, and the degree of hydrolysis after exposure is determined by measuring the change in the optical rotation of the solution as the sucrose hydrolyses to invert sugar.
An integrated temperature or "ecological mean temperature" T e (Berthet, 1960) is calculated from the change in rotation. This is thought to be a more meaningful figure in the context of ecological studies than the standard mean temperature, because the temperature quotient (Q10 ) of the reaction is 4.8 (Lee, 1969) and thus in the range of that of many life processes (2 to 6). Lee points out, however, that simply because this reaction and biological reactions have exponential temperature functions, a simple relationship between those functions does not necessarily exist. Still, the sucrose inversion method appears to be a better indicator of integrated biological activity than the standard mean (Berthet, 1960), as well as being cheap and ideal for use 187 in an area remote from weather stations. It has been used in this study for those reasons.
Test tubes were filled with 20 mL of the buffered sucrose solution (the "fast solution" . of Berthet, 1960) and kept at -20 °C until needed. Two tubes were placed at the soil-litter interface at Anglers Creek monthly or more frequently and a control tube was taken into the field, thawed with the field tube and returned to the freezer, to account for inversion occurring during transport. Exposed tubes were refrozen until analysed.
Measurement of the optical rotation of the sucrose solutions was performed using a Bellingham and Stanley Model D2 polarimeter. Te was calculated for each interval of exposure and these temperatures are shown in Figure 5.5, with maximum and minimum readings for the same periods.
As population sampling extended for longer than the period covered using the sucrose inversion method, it was necessary to estimate T based e on Bureau of Meteorology records of temperatures at Lake Leake Chalet, and the correlation between these and measured T . The Bureau records monthly e averages of daily maximum and Minimum temperature; however the ecological means were measured over different intervals. Each day of the year was therefore assigned the average for that month as recorded by the Bureau, and means were calculated for the periods over which sucrose solutions were exposed. The relationship between these values and T e from
Anglers Creek was surprisingly close: of the functions tried, the best fit was the linear relationship with the averaged maximum values from
Lake Leake, which gave a correlation coefficient of 0.97. This relation- ship was found to be (n = 17):
T e = -1.03 + 0.67 X where X = averaged maximum temperature at Lake Leake Chalet (see above)
T = ecological mean temperature at e Anglers Creek. 22— 20— 18—
, 16—
cu =r-12— .4--
atrI2) 10
8 — 6— 4— 2— 0—
11 1 1 1 1 1 1 1 A SO N DJ F M A NIJ J A 1976 1977
Figure 5.5 Maximum, minimum and ecological mean (T e ) temperatures at soil/litter interface at Anglers Creek site, over various intervals during period of study. 188 and is plotted in Figure 5.6. Using this formula, it is possible to calculate T for any period for which there are Bureau records, as the e data used to derive the relationship applied to time intervals throughout the year. Monthly values Of T e during the population study (Chapter 8) were calculated and appear in Figure 8.4.
Section 5.4 Relative Humidity Measurement
As landhoppers are extremely susceptible to desiccation, it is reasonable to expect that the water content of the air is a rather important feature of their environment. Relative humidity was therefore monitored by the use of a Casella Thermo-hygrograph, which was placed on the forest floor in the study area, under a shelter consisting of a 60 cm x 60 cm sheet of polythene held 60 cm above the litter by four posts. This arrangement was preferred to the usual louvred box because it allowed the hair hygrometer to be closer to the litter surface. In fact it was
8-10 am above ground level. The shelter was moved every three months to counter the long-term effects of sheltering the forest floor from rainfall and litterfall.
The thermo-hygrograph has a clockwork mechanism and runs for about ten days between windings. As Anglers Creek was visited only monthly or fortnightly during the sampling period, only part of the year was covered. The seasonal changes in the daily relative humidity cycle was shown quite clearly in the record of the 170 days over 13 months on which measurements were made.
Chart records relating to several days in summer and winter are shown in Figure 5.7. The humidity trace at 100% Was not always aligned with the graduation of the paper. The plateau level, rather than the
100% graduation was therefore used in calibrating the trace. The scale of the pen movement was found to correspond to the chart scale.
The proportion of time for which the relative humidity fell within arbitrary limits was calculated. These limits were 50-60%, 60-70%, — 12-
5 10 15 20 25 Average of mean maximums, Lake Leake
Figure 5.6 Relationship between ecological mean temperature and average of mean maximum temperatures (see text for explanation) at Anglers Creek study site. r = correlation coefficient. Monday Tuesday Wednesday I Thursday Friday Saturday Sunday 810 2 46 8 . 10X 6 8 1 112 4 6 8 10 112 4 68 1 12 4 6 8 10XII 2 4 6 8 10 12 4 6 8 10 2 4 6 8 102(112 4 6 8 10 2 4 6 8 10 6.. 8 10 12 4 66 fehtiir 111111=EITIONC .,_•; 120 fiDni 0. TIONINEMINI 9 0 VAII11111111188111 lFi N=45165
M _7 0 - 1674111116120 Illa°141° 60 60 60 : 0 111111111111116411. 50 50 56'
Nig 4 111111,1111111111 40 40
b=1- 11111111111Mili30 30 r PR' D INt BIWA20 INSIMEISS 20 • Monday Tuesday Wednesday Thursday Friday Saturday Sunday 810 2 4 6 8 10 12 4 6 8 10X112 4681 12 4 6 8 10 112 4 68 10X 2 468 10XII 2 4 681 12 4 6 8 10 1 2 4 6 8 10 12 4 6 8 10X1I2 4 6 8 10X I 2 46 8 10 I 2 4 6 8 10Xo2 46 8 20 120 .120.. . If24INEF &lam, 110 — NUS& UMW! ItIWINN 100 100 WOMM 011 MINIMA • . wing 90 90 90 I. — 80 80 LU 70 0
60 60
50 Cr • 40
ML 111111111=111 30 .30 Inst..No. D \IN \ Year.77.-i.Monthli 20 20
Figure 5.7 Copy of temperature-humidity recorder charts for 10-day periods in February and July 1977 at Anglers Creek study site 189
70-80%, 80-90%, 90-95%, above 95% and below 100%, and on 100%.
Figure 5.8 shows how the time covered by hygrograph records was
divided between these values, and also shows the daily rainfall as
recorded by the Bureau at Tooms Lake.
These R.H. measurements reveal several points of interest.
Firstly, the seasonal variation is well marked and is related to temperature
(Figure 8.4) as well as rainfall. Secondly, the general level of R.H.,
even during dry periods, is high, never falling below 50%, and most of the year, remaining above 95%. Thirdly, the diurnal cycle of relative humidity
on the forest floor is extremely marked in the drier months, always return-
ing to high values overnight. This could have important implications with
respect to the diurnal activity cycle of the amphipod species present
[see Section 7.2(b)]. On dry days, the R.H. minimum occurs at about the
same time as the temperature maximum (usually around mid-afternoon) while
the R.H. maximum on these days begins at or before the temperature minimum, near dawn.
Section 5.5 Botanical description
The site is in low closed-forest (Specht, 1970) with coppiced
Musk (Olearia argophylla F. Muell.) providing 100% canopy cover, with
several emergent Stringy-bark Gums (Eucalyptus obligua L'Herit.) and
Black Wattles (Acacia mearnsii De Wild.). Apart from occasional
Soft Treeferns (Dicksonia antarctica Labill.) most of the area is open under the trees. The openness of the forest floor is interrupted by several logs and patches of Mother Shield-fern (Polystichum proliferum
Presl.) under the trees.
Where the canopy is broken, several small Sassafras trees
(Atherosperma moschatum Labill.), Stinkwood (Zieria arborescens Sims.),
Cheesewood (Pittosporum bicolor Hook.) and Coprosma nitida Hook. f. are found, with Batswing Fern [Histiopteris incisa (Thumb.) J. Sm.], Ruddy
Ground-fern [Hypolepis rugosula (Labill.) J. Sm.] and Austral Bracken 1977 38 -
36 - 34 -
32 -
30 - 28
26 - 24 -
22 -
12 - 10 -
8 -
6 -
2 - El,. ..II ILL I it ) I OCT OV DE JAN FEB MAR APRIL MAY JUNE JULY AUG SEPT Relative humidity 11 100% 9095% 70 - 80% 50-60% values HIM
95-100% NUNN 80-90% 60 -70°/o 1•0 T EMI 9 II.
8
7 -0
0 .6
•_ 5 II1 0 0
0
•3
2
DEC JAN FEB MAR APR MAY JUNE JULY AUG SEPT OCT NOV DEC 1976 1977
Figure 5.8 Proportion of time monitored when relative humidity fell in selected intervals, near forest floor at Anglers Creek study site. Each column represents one 10-day period during which the recorder ran. Daily rainfall at Tooms Lake in 1977 also shown. 190
[Pteridium esculentum (Forst.) Diels] in thick stands.
• Plate 5.3 gives a view of the forest in which the study area is located. The tall E. obligua and smaller A. mearnsii stand Above the closed 0. argophylla canopy. Plate 5.4 shows part of the study site in the forest; most of the area is more open than this.
Much of the site, including the section used for population sampling (Figure 7.2) is rather homogeneous, with the forest floor open under the Olearia canopy. This apparent homogeneity, as well as the ease of movement and high amphipod density, was amongst the qualifications of the site for this study.
The litter layer varies in depth between 2 cm and 5 am during the year, and is composed largely of Olearia leaves (which are 10-15 am long: Plate 5.5) with Olearia bark and twigs, and the litter of Acacia and Eucalyptus in smaller quantities. The fermentation layer is 1-2 cm thick and is composed largely of amphipod faeces, mixed with Acacia twigs and at times Acacia and Olearia inflorescences. Below an indefinite humus layer, brown clayey loam extends for at least a metre further
(Plate 5.6). Near the soil surface, there is a concentration of Olearia
roots and rootlets.
Section 5.6 Phenology
During the course of the year, several events have noticeable
effects on the study site. In summer, the maximum fall of Glearia and
Eucalyptus leaves occurs (see Section 6.3). This causes a marked increase
in the thickness of the litter layer, which is at its thinnest in late
spring. The Acacia trees flower from October to December, when dead
inflorescences are conspicuous in the litter. In November and December,
the Olearia is in flower, and new fern growth is evident at this time and
through summer. New light green fronds appear on Polystichum, Dicksonia
and Pteridium, while Histiopteris, which dies off in winter, sends up
copious new growth in the more open areas near the creek. E. obLigua Plate 5.3 Forest at study site, Anglers Creek, Treeferns Plate 5.4 Inside the forest near the study site (Dicksonia) line creek just beyond road margin. at Anglers Creek. Light green foliage is Olearia argophylla. Eucalyptus obliqua (dark trunks)and E. viminalis (light trunks) stand in background. Plate 5.5 Newly-fallen Olearia argophylla leaves, shown on millimetre graph paper.
Plate 5.6 Soil profile (pit 1 m deep) at Anglers Creek study site. 191 flowers in April and spent stamens appear in the litter fall. Over winter, the litter on the forest floor is entwined with basidiomycete rhizomorphs similar to those of Clavariadelphus junceus (Fr..) Corner described by Ashton (1975) in E. regnans forests in Victoria.
Section 5.7 Soil and litter characteristics
The water content of the litter and of the soil, as well as the weight loss of the soil on ignition, were determined during the study.
Methods and Results
On four occasions, 4 January and 9 February 1977, 25 May 1978 and 1 December 1979, samples of the forest floor were taken for measure- ment of their dry weight and, except on 25 May 1978, their water content.
Two samples on each of the first two dates, three samples on the third and ten on the fourth date were collected by removing all litter down to the compacted soil (i.e. litter and fermentation layers were removed) inside an open-ended steel cylinder of 32 cm diameter. This cylinder was placed on the litter surface and the litter was cut around its lower edge with a sharp kRife. In the laboratory, the litter was weighed on a
Mettler P1210 balance, dried for 24 h at 105 °C, then reweighed. The standing Crop of litter on these dates is shown in Table 6.1, while the litter water content, where measured, appears in Table 5.1.
As outlined in Section 8.2, twelve samples of soil and litter were collected monthly for amphipod population assessment. These samples
7 cm in diameter, and containing all surface litter and soil to a depth of 7 cm, were weighed on return to the laboratory and after the extraction o of animals, the weight lost on drying for 24 h at 105 C was determined.
The percentage water content of these samples gives a good indication of the soil water content, as they contain much more soil than litter.
Table 5.2 shows the water contents thus determined. 192
TABLE 5.1 Water content of litter samples from study site.
Date % water
4 January 1977 59..7 ± 10.2
9 February 1977 36.1 ± 0.6
1 December 1979 38.9 ± 10.6
TABLE 5.2 Water content of soil/litter samples from study site.
Date % water + S.D.
11.viii.1976 53.7 1
8.ix.1976 55.6 1
8.x.1976 57.7 1
10.xi.1976 57.2 1
8.xii.1976 not measured
4.i.1977 58.0 1
9.ii.1977 46.2 ± 2.4 10
9.iii.1977 49.1 ± 5.1 12
14.iv.1977 48.6 ± 4.1 12
10.v.1977 55.2 ± 6.0 12
8.vi.1977 55.5 ± 6.1 12
8.vii.1977 53.8 ± 6.6 12
17.viii.1977 53.9 ± 5.4 12
19.ix.1977 56.1 ± 5.1 12
26.x.1977 52.2 ± 5.2 11
26.x.1977 (2nd site) 39.3 ± 4.0 12
2.xii.1977 51.5 ± 7.0 12 193
An indication of the organic content of soil from the Anglers
Creek area was gained by ashing a sample of soil, taken from below the
fermentation layer. 3.31 g of soil placed in a muffle furnace (Scien-
tific Equipment Manufacturers) at 500 °C for 48 h yielded 1.37 g of ash
corresponding to a "weight loss on ignition" of the soil of 58.6%.
Discussion
From the first two determinations of litter water content, the
loss of water from the litter layer in even a short period of drought
is demonstrated.
The first two determinations of litter water content demonstrate
the loss of water content from litter layer in even a short period of
drought. Rain fell on January 3 and had fallen each day from 23 to
28 December inclusive (Figure 5.3). Before 9 February, however, no
appreciable falls were recorded for 16 days, allowing a large drop in
the moisture content of the litter.
The soil, on the other hand, holds the moisture much more securely.
The soil/litter water content is fairly stable throughout the year and
the mean values display a total range of less than 10%. The lowest
values, as might be expected, occur in late summer and early autumn, when
the evapotranspiration rate is highest. The high organic content of the
soil indicated by the "weight loss on ignition" of 58.6% is largely respon-
sible, together with its friable, open structure, for this high water
retention. This, in turn, must ensure that the relative humidity at the
soil/litter interface, despite the fluctuations of the litter moisture
content, remains at 100%.
Section 5.8 Fauna
No detailed investigation of the vertebrate fauna of the site
was made, but a number of species were sighted during visits to the
area. As most of these visits were day trips from Hobart, two hours 194 drive away, the best times for observation were missed. However, species observed in the area are recorded below.
Mammals
Bennett's Wallaby Macropus rufogriseus
Dusky Marsupial-mouse Antechinus swainsonii
Brush-tailed possum Trichosurus vulpecula
Cdroppings only)
Birds
Scarlet Robin Petroica multicolor
Grey Fantail Rhipidura fuliginosa
Superb Fairy-wren Malurus cyaneus
Green Rosella Platycercus caledonicus
Black Currawong Strepera fuliginosa
Forest Raven Corvus tasmanicus
Yellow-tailed Black
Cockatoo Cal yptorhynchus funereus
Reptiles
Tasmanian Tiger Snake Notechis ater
Possible predators of amphipods
Antechinus swainsonii was observed turning dead leaves over as if in search of food. This was at midday, an unusual time for this timid nocturnal animal to be feeding; however this observation indicates that this species might feed on amphipods, which are by far the most common macro-invertebrates present.
Of the birds seen, the Scarlet Robin, the Grey Fantail and the
Superb Fairy-wren are the most likely to be predators of amphipods, although they were not seen under the Olearia canopy, but out in the open area near the road. However, at no time were birds seen turning over litter in the 195 study area. It appears that there are no major vertebrate predators of amphipods present, whose activities might be expected to have a significant effect on the population.
Invertebrates
Extraction of invertebrates from litter and soil taken from the study site over 1977, both by hand and using modified Tullgren apparatus yielded representatives of a number of groups which are listed below. 196
Ph. Nematomorpha
Cl. Gordioidea
Ph. Annelida
Cl. Oligochaeta
Ph. Mollusca
Cl. Gastropoda
0. Pulmonata
Ph. Arthropoda
Cl. Arachnida
O. Pseudoscorpiones
O. Araneae
O. Opiliones
O. Acarina
S-o Mesostigmata
S-o Trombidiformes
S-o Sarcoptiformes
Cl. Crustacea
O. Isopoda
S-o Oniscoidea
O. Amphipods
S-o Gammaridea
Super-f. Talitroidea
F. Talitridae
Mysticotalitrus tasmaniae
Mysticotalitrus crypticus
Keratroides vulgaris
Keratroides angulosus
Cl. & O. Collembola
S-o Arthropleona
Super-f. Poduroidea 197
Super-f. Entomobryoidea
S-o Symphypleona
F. Sminthuridae
Cl. Insecta
0. Plecoptera
F. Grypopterygidae
Dinitoperla opposita (Walker)
O. Hemiptera
S-o Hamoptera
Super-f. Fulgoroidea
S-o Heteroptera
Super-f. Dipsocoroidea
F. Dipsocoridae
Super-f. Lygaeoidea
F. Lygaeidae
S-f. Rhyparochrominae
Other Lygaeoidea
Other Heteroptera
O. Coleoptera
S-o Adephaga
Super-f. Caraboidea
f. Carabidae
S-o Polyphaga
Super-f. Staphylinoidea
F. Scydmaenidae
Other Staphylinoidea
Super-f. Scardbaeoidea
F. Scarabidae
Super-f. Dryopoidea
Super-f. Elateroidea
F. Elateridae 198
Super-f. Cucujoidea
F. Corylophidae
F. Tenebrionidae
F. Lagriidae
F. Melandryidae
Super-f. Chrysomeloidea
F.• Chrysomelidae
Super-f. Curculionoidea
F. Curculionidae
0. Trichoptera
Super-f. Rhyacophiloidea
F. Calocidae
Caloca saneva (Mosley)
see Neboiss (1979)
0. Lepidoptera
O. Diptera
S-o Nematocera
F. Psychodidae
F. Chironomidae
Other Diptera
Cl. Diplopoda
Cl. Pauropoda
Cl. Chilopoda
O. Geophilomorpha
Cormocephalus westwoodi 199
None of these groups represented a biomass approaching that of the landhoppers. Trichopteran larval cases were very conspicuous and common, but many of these were empty, as they lasted quite a long time after the adults had emerged. Carnivores large enough to tackle amphipods included carabid beetles, mygalomorph spiders and chilopods: by far the most common of these was the centipede Cormocephlus westwoodi which grows to about 60 mm and might be a significant predator of amphipods. 200
CHAPTER SIX
LITTER FALL AND DECOMPOSITION
A study of organic input to the forest floor at the
Anglers Creek study site
Section 6.1 Introduction
Forest litter consists of dead or dying leaves, wood, flowers. , fruits, seeds, bark and other debris shed by plants, thus forming a downward flow of material in the forest ecosystem. The fall of litter, in fact, is a major pathway for both energy and nutrient transfer in forests (Bray and Gorham, 1964).
A large proportion of the nutrient removed as litter is stored in the litter and soil strata. The release of part of this nutrient pool to allow its return to the photosynthetic machinery is fuelled by the energy removed from it as litter. Transfer of nutrients from the plants to the soil through litter fall occurs in two stages; the initial fall from the canopy to the forest floor, and the subsequent decomposition of the litter (Attiwill, 1968). Leaching is important, especially for the more soluble nutrients, in each of these stages.
The accumulated litter forms both habitat and energy source for a host of life-forms. These may use the litter directly as an energy source (detritivores and saprophytes on litter), or they may use it indirectly (by predation, microfloral grazing and breakdown of dead fungal and animal material).
The soil/litter fauna has been seen as a regulator of the decom- position segment of the forest ecosystem (Crossley, 1977), and the ultimate aim of this investigation is to determine the role of land amphipods (as detritivores) in this regulation at Anglers Creek.
Knowledge of the dynamics of litter fall and accumulation is obviously important in understanding the role of decomposers, and for this 201
•reason a detailed study of litter fall was carried out. This is reported in the following Chapter.
Section 6.2 Methods
Litter fall
Measurements of the rate of litter fall at Anglers Creek were made from early August 1976 to early December, 1977. Three square bins of side 620 mm, similar in design to those used by many workers, including
Ashton (1975), Golley et a/. (1975) and Rogers and Westman (1977), were placed in different parts of the population sampling site (Figure 7.2).
These bins consisted of a wooden frame 14 am high, with a floor of gal- vanized flywire (44 meshes per 10 cm) which sat on another frame 7 cm high, so that the mesh was held off the forest floor. A muslin sheet was placed across the bin and attached with string stretched around the upper frame.
Each month the litter was collected by removing the muslin sheet with the litter inside it, cutting off any projecting pieces of wood or bark
level with the side of the frame. The muslin was taken to the laboratory and
dried at 105°C. The dry litter, which contained a significant amount of
small material, was then easily removed from the muslin. A Mettler HlOW
analytical balance was used to weigh the litter after it was sorted into
the categories shown below. Dried litter was stored at room temperature
for up to a year before analysis for nutrient content.
Litter categories:
Olearia leaves
Eucalyptus leaves
Other leaves
Olearia bark
Eucalyptus bark
Acacia bark
Branches, twigs and nuts (mood above 2 mm diameter) Small litter 202
During annual leaf-fall, the bipinnate leaf of Acacia mearnsii first drops its leaflets, then the rachis loses the remains of the pinnae and falls itself. The 2 mm diameter chosen to define "branches and twigs" distinguished between the larger twigs of Olearis and Acacia rachides included in "Small litter". The latter category thus contained
Acacia leaf components, Olearia and Acacia flowers, Eucalyptus stamens, herbivore frass, tiny bark and leaf fragments and wind-blown dust.
Only three bins were used to measure litter fall; although a greater number would have been desirable, it was decided that as the study of litter fall was not the major emphasis of the project, the extra work involved would not yield a proportionate return. Many workers have used 10 litter bins (e.g. Golley et al., 1975; Christiansen, 1975;
Rogers and Westman, 1977) although Newbould (1967) has recommended the use of a minimum of 20 bins. In this case, the small area of the vegetational type under study, the homogeneity of the Olearia canopy, and the lack of large amounts of branch material (which is highly aggreg- ated, spatially and temporally) all indicate that a reasonably accurate estimate is possible from a small number of bins.
Standing crop of litter
The total weight of forest floor litter was determined on several occasions, and the method is outlined in Section 5.7. The location of each sample was selected more or less at random, but only in open patches of forest floor. The percentage of the sampled area which comprised open forest floor was determined by measurements on three randomly-chosen 20 m transects. As stated before, forest floor litter was collected on four occasions: 4 January 1977, 9 February 1977, 29 May 1978 and 1 December 1979.
Nutrient analysis
Analysis of the nitrogen, phosphorus, potassium, calcium and copper content of litter from bins, litter from the forest floor and soil from the top 10 am was performed by the Government Analyst Laboratory, 203
Hobart. Separate analyses were made of Olearia leaves, bark and
twigs, Eucalyptus leaves, bark, twigs and nuts, Acacia rachides,
Dicksonia fronds, and small litter. Most of the newly-fallen litter
analysed was collected in January and February 1977. Soil was collected
below the fermentation layer, to a depth of 5 cm. Forest floor litter
was divided into upper litter, with large leaves, twigs and bark and
lower litter, incorporating the fermentation layer (amphipod faeces)
but including Acacia leaflets and rachides. Soil and litter were
collected for nutrient analysis in 1 December 1978. Forest litter and
soil samples were sorted through manually, and all rocks and as many
animals as possible, removed. Five samples of each of these litter and
. soil categories were analysed, except for eucalypt nuts (two samples) and
Acacia rachides (three samples).
The proportions of upper and of lower litter present in the
litter layer were determined from forest floor samples collected on
1 December 1979, when the fraction of the litter layer comprising leaves
was also measured. Soil samples were collected at the same time so that
the dry weight of soil down to 5 cm could also be found.
The analysis techniques used were as follows:
Nitrogen: Kjeldahl digestion
Phosphorus: Wet ashing with nitric acid/perdhloric acid,
' then colorimetric analysis with vanado-molybdate
reagent.
Calcium: -- Wet ashing with nitric acid/perchloric acid then
Potassium: analysis by atomic absorption.
Copper:
Section 6.3 Results
As litter fall was collected for sixteen months, an annual mean
was calculated by averaging the litter fall in the first twelve and the
last twelve months. 204
The mean annual litter fall was 9390 kg/ha. Of this,
3344 kg/ha, or 35.6% consisted of large leaves (mainly Olearia and
Eucalyptus) while 19.2% was wood and 6.1% bark. The remainder (39.1%) was "Small litter", the composition of which Changed over the year.
Over summer, Olearia and Acacia flowers were important, while at the same time the total amount of Acacia leaflets was at its highest.
Figures 6.1 to 6.4 show the monthly distribution of the fall of total litter, wood and bark, Olearia leaves, Eucalyptus leaves and small litter over the sampling time. Daily means were calculated for each collection period, and from these, monthly totals computed.
The annual periodicity of total litter fall is due to summer peaks of leaf fall, which is also expressed in the peak of small litter.
During this peak, the litter bin which was beneath an Acacia tree always contained the highest amount of "small litter", in contrast to the rest of the year, when the distribution of "small litter" between the bins was more even. This indicates the importance of Acacia litter components in contributing to the "smell litter" peak, which occurs in March, compared with the January maximum of both Olearia and Eucalyptus leaf fall.
November leaf fall values (especially Olearia) were noticeably higher in 1977 than in 1976. Comparison of the calculated ecological mean temperatures for the spring months of each year (Figure 8.4) indicates that the earlier rise in temperature in 1977 caused the advance of the summer growth peak, accounting for the earlier fall of leaves.
• Twig and bark fall, on the other hand, did not vary consistently with temperature over the sampling period, nor did it appear to be related to the amount of wind in each month, as measured by the total distance run
(Met. Bureau records, Lake Leake Chalet).
The results of litter standing crop measurement were corrected for the percentage of forest floor obstructed by trees and logs, on which litter does not build up. Knowledge of this figure (11.6 11.3%)- allows 1976 1977 6000- w 5000- 4000- a), -Jm 3000- Er 2000 - .C7 C 1000 -
SI/
500-
Figure 6.1 Distance run monthly by wind at Lake Leake Chalet during period of study (Bureau of Meteorology records), and monthly litter fall at Anglers Creek study site. Wood and bark component of litter fall represented by solid area.
Figure 6.2 800 700 600 Small litter 500 400 2 300 : 200 _c 100
300 Figure 6.3 200 Eucalyptus leaves 100 1 Figure 6.4 400 Olearia leaves 300 200 100
AS 0 N D J F MAMJJ ASO N 1976 1977
Figures 6.2-4 Monthly fall of small litter, eucalypt leaves and Olearia leaves at Anglers Creek study site during period of study. 205
TABLE 6.1 Litter standing crop at Anglers Creek, corrected for
area obstructed by trees and logs.
Date Litter standing No. of crop (kg/ha) measurements
4 January 1977 6540 2
9 February 1977 8134 2
29 May 1978 14139 3
1 December 1979 9513 10
1 December 1979, leaves only 2447 10
TABLE 6.2 Nutrient analysis of litter and soil from Anglers Creek.
Items 1-10 from litter bins in 1977, items 11-13 collected
1 December 1978.
Material. %N %K %Ca ppm Cu
1. Olearia leaves 0.91±0.09 0.52±0.06 1.12±0.06 0.50±0.06 5.80±0.33 5
2. Olearia bark 1,02±0.13 0.11±0.04 0.64±0.09 0.50±0.13 8.86±1.28 5
3. Olearia twigs 0.76±0.06 0.31±0.25 0.47±0.11 0.50±0.03 8.88±3.14 5
4. E. obliqua leaves 0.72±0.08 0.16±0.01 0.56±0.02 0.69±0.12 4.42±0.91 5
5. E. obliqua bark 0.34±0.02 0.10±0.01 0.18±0.08 0.17±0.04 6.06±2.00 5
6. E. obliqua twigs 1.05±0.28 0.15±0.03 0.52±0.10 0.38±0.13 6.58±1.47 5
7. E. obliqua nuts 0.47±0.00 0.17±0.04 0.23±0.02 0.33±0.01 3.90±0.18 2
8. Acacia twigs 1.87±0.14 0.38±0.06 0.40±0.03 0.65±0.12 7.07±0.12 3
9. Dicksonia fronds 1.97±0.07 0.18±0.03 0.39±0.04 0.40±0.07 10.56±2.31 5
10. Small litter 2.12±0.02 0.35±0.05 0.54±0 .02 1.01±0.03 11.60±0.62 5
11. Soil, 0-3 cm 1.14±0.15 0.13±0.02 0.21±0 .07 3.99±0.29 61.80±10.38 5
12. Lower litter 1.23±0.03 0.12±0.00 0.50±0 .23 2.62±1.10 37.24±18.32 5
13. Upper litter 0.97±0.06 0.16±0.00 0.59±0 .04 0.74±0.16 9.52±2.41 5 206 comparison of the standing crop with the rate of litter fall on an areal basis. The average litter standing crop on each of the four dates on which it was measured is shown in Table 6.1. On the basis of the variation of litter fall throughout the year (Figure 6.1), the standing crop of litter might be expected to increase steadily through
summer, reaching a peak in autumn. Taking year-to-year differences
into account, this pattern is basically demonstrated by the data in
Table 6.1. If the forest floor accumulation is in equilibrium with
litter fall, the pattern for winter and spring will be a similar decrease,
so that an approximate annual mean standing crop could be estimated by
averaging the four measurements in Table 6.1, as they cover the summer-
autumn period. This mean forest floor weight is 9582 kg/ha.
Many workers in the United States and Australia (Kittredge, 1948;
Jenny et al., 1949; Olson, 1963; Galley et al., 1975; Ashton, 1975;
Rogers and Westman, 1977, etc.) have used the ratio of annual litter fall
to standing crop of litter as a measure of litter decomposition rate.
Olson (1963) used this ratio to calculate the half-life of the annual
litter fall in a forest where equilibrium of the litter accumulation has
been achieved. This ratio, k, for the study site is 0.980 and the
litter half-life is 0.707 years, while for leaf material only, these values
are 1.367 and 0.507 years respectively.
. Another indication of the rate of disappearance of the leaf com-
ponent of the forest litter was provided by observation of the forest
floor under the litter traps. These traps were placed in early August
1976, and by March 1977 most leaves had gone, and the few remaining (about 2 50/m or 150 (kg/ha) showed signs of attack. Most of the leaves under
the bin in August would have reached the forest floor during the leaf fall
peak of early 1976, so this leaf litter lasted about one year. Where
there is no litter input, as in this experimental situation, the tail of
the exponential decomposition curve (Jenny et al., 1949) will be cut off 207 by the activities of the resident detritivores, as food becomes
scarce. The real life of the leaf litter would thus be rather more
than a year, corresponding to the estimated half-life of about six months for leaves, from the forest floor ratio, k.
The litter layer in the forest on 1 December 1979 was found to be 25.7% leaf material, and the "lower litter" category made up 18.1%
of the standing crop. The dry weight of soil down to 5 cm depth on 2 an areal basis was 24.7 kg/m .
The results of nutrient analysis are shown in Table 6.2, while
Table 6.3 gives a comparison of some of these concentrations with those
found by workers in other parts of Australia. Where comparison is possible, the values obtained are in good agreement with those of Ashton
(1975) in Victorian Eucalyptus regnans forest. 0. argophylla leaf
nutrient contents are similar, and the "small litter" values, except for phosphorus content, are close to Ashton's Acacia dealbata (a species
closely related to A. mearnsii) leaf analysis results. With respect to
E. oblique litter, the content of nitrogen, potassium and calcium is in
the same range as other eucalypts where data are available (E. marginate -
Hatch, 1955; E. oblique - Attiwill, 1968; E. regnans - Ashton, 1975;
E. signata and E. umbra - Rogers and Westman, 1977).
The phosphorus content of the E. oblique leaf litter at Anglers
Creek, however, is 2-3 times greater than that of all the other eucalypt
species mentioned except E. umbra, from North Stradbroke I., Queensland, which contains slightly more phosphorus (Rogers and Westman, 1977).
This element is also much higher in "small litter" from Anglers Creek
than in A. dealbata from Victoria.
Amongst the studies listed in Table 6.3, copper content was measured
in only one (Rogers and Westman, 1977) and the levels found in the leaf
litter of eucalypts and understorey species are 2-5 times as high as those in the Anglers Creek leaf litter, although the levels in wood and bark are very similar. TABLE 6.3 Nutrient content of selected litter components in a range of forests
Component Species Place $N %K %Ca ppmCu Reference
Leaf E. marginate Dwellingup, W.A. 0.47 0.25 0.77 0.22 Hatch, 1955 Litter E. regnans Hume Plateau, Vic. 0.87 0.12 0.55 0.30 Ashton, 1975 E. signata N. Stradbroke I. Qld. 1.07 0.14 0.60 0.39 17 Rogers & Westman, 1977 E. umbra N. Stradbroke I. Qld. 0.87 0.11 0.73 0.83 18 Rogers & Westman, 1977 E. oblique Anglers Ck.,Tas. 0.72 0.16 0.56 0.69 4.4 This study 0. argophylla Hume Plateau, Vic. 1.20 0.21 0.83 0.50 Ashton, 1975 0. argophylla Anglers Ck., Tas. 0.91 0.52 1.12 0.50 5.8 This study A. dealbata Hume Plateau, Vic. 2.03 0.24 0.63 0.55 Ashton, 1975
Small litter (A. mearnsii) Anglers Ck., Tas. - 2.14 0.35 0.54 1.01 11.6 This study Leaf litter Tristania N. Stradbroke I. Qld. 14 Rogers &Westman, 1977 Angophora N. Stradbroke I. Qld. 9 Rogers & Westman, 1977 Persoonia N. Stradbroke I. Qld. 22 Rogers & Westman, 1977 Pinus strobus S. New Hampshire, U.S.A. 0.45 0.20 0.65 0.50 Kittredge, 1948 Acer rubrum S. New Hampshire, U.S.A. 0.60 0.53 1.20 1.90 Kittredge, 1948 Leaves & wood E. oblique Hume Plateau, Vic. 0.09 0.51 0.18 Attiwill, 1968 Twigs & bark E. marginata Dwellingup, W.A. 0.38 0.20 0.95 0.18 Hatch, 1955 Forest floor E. regnans ' Florentine Valley, Tas. 0.12 0.71 0.53 Harwood & Jackson, 1975 litter Nothofagus Atherosperme Acacia E. oblique Anglers Ck., Tas. 0.97 0.16 0.59 0.74 This study 0. argophylla A. mearnsii 208
Section 6.4 Discussion
The annual litter fall at the Anglers Creek site is high, compared with other Australian eucalypt-dominated formations (see
Table 6.4). This is partly due to the 100% canopy cover afforded by
the understorey of Olearia: however it probably also indicates a par-
ticularly fertile site (Bray and Gorham, 1964).
Much of the literature about litter fall concerns northern
deciduous forests (Bray and Gorham, 1964), in which a very large proportion
of the annual fall of leaves occurs in autumn, as temperatures fall.
Although eucalypts are not deciduous, they also have a very marked peak
of leaf-fall, but this is in spring and early summer (Hatch, 1955; McColl,
1966; Webb et al., 1969; Ashton, 1975; Rogers and Westman, 1977) with the
new growth as temperatures rise. Slightly different timing occurs in
other Australian forests: while earlier maximum leaf-fall is found in
subtropical rainforest (Webb et al. 1969), Nothofagus-dominated temperate
rainforest displays an early autumn peak (Howard, 1973).
The annual progression of changing environmental conditions for
litter decomposers is thus different in eucalypt forests from that exper-
ienced in deciduous forests. The litter layer is thinnest in spring and
thickest in summer, so when the forest floor is most likely to dry out,
the extra layer of leaves helps retard the loss of moisture, reducing the
chance of desiccation of the cryptozoa:
Initial decomposition of eucalypt leaves occurs under the influence
of high temperatures, so if the litter does not become excessively dry,
fungal attack might be expected to proceed quickly. By autumn, fungal
colonization, leaching in wetter areas and relatively warm climate would
tend to encourage the activity of the detritivores. The highest rate of
leaf decomposition recorded in Australia is in warm temperate eucalypt
forest in New England, where the climate is warm in summer, while rainfall
is high and occurs throughout the year (Richards, 1976 - see Table 6.5). TABLE 6.4 Rate of litter fall and standing crop of litter (dry weight) in some Australian forests
Locality Dominant Stand Annual Litter Reference Species Type litter fall standing crop kg/ha/y kg/ha
Southwest W.A. Eucaluptus diversicolor Mature 5650 Stoate, 1958 Regrowth 6030
Ddellingup, W.A. E. marginate Mature 2377 18930 Hatch, 1955 Pole 3097 (22 y since Sapling 2575 last fire)
Dwellingup, W.A. E. marginata Mature 9200 O'Connell, Grove & Dimmock,
forest, 10700 1978 different 15000 soil types 16900 6 y since 18400 cool fire 13000 13200
Mt. Lofty Ra., S.A. E. oblique & 2330 9800 Lee & Correll, 1978 E. baxteri
N. Stradbroke I., E. signata 6426 27000 Rogers & Westman, 1977 Qld.
Nr. Brisbane, Qld. E. umbra, 3108 10237 Birk, 1979 E. bailegana, including including Angophora woodsiane grass grass
Whian Whian N.S.W. E. pilularis 6507 Webb et al., 1969 (leaves only)
Taree, N.S.W. E. pilularis 25 y regrowth 4325 15357 van Loon, 1969 24 y regrowth 7088 39 y regrowth 5594
Benandra, N.S.W. E. maculate Mature 6846 McColl, 1965 Pole 5063 Sapling 3601
Locality Dominant Forest Annual Litter Reference
Species type litter fall standing crop kg/ha/y kg/ha
Sydney, N.S.W. E. piperita 508 6280 Hannon, 1956, 1958 E. pilularis 384
Hume Plateau, Vic. E. obliqua Mature 3560 18250 Attiwill, 1968
Hume Plateau, Vic. E. regnans Mature 7756 22250 Ashton, 1975 Spar 7731 Pole 6606 20250
N.W. Tasmania E. obliqua Mature 7100 40800 Madden et al., 1976 50 y regrowth 6700 63200 20 y regrowth 4400 42000 13 y regrowth 3500 32500 5 y regrowth 3500 10200
Florentine Valley, E. regnans, 40100 Harwood & Jackson, 1975 Tas. Nothofagus, Atherosperma, Acacia
Anglers Creek, E. obliqua, 9390 9582 Present study E. Tas. acacia, Olearia 209
Clark's findings (1954) that leaves lay for at least one to two months before attack by amphipods were substantiated at Anglers Creek: other workers (e.g. Satchell and Lowe, 1967) have made similar observations on other cryptozoan groups. Palatability of leaves is apparently in- creased by leaching and fungal attack. Food for amphipods is probably plentiful from summer through to early spring, when the litter layer becomes noticeably thinner.
The standing crop of litter at Anglers Creek is smaller than
that in many other eucalypt forests, as shown in Table 6.3, which gives
the weight of accumulated litter (forest floor) found in a number of
Australian studies.
Use of the ratio of annual litter fall to litter standing crop
to predict time functions of litter decomposition must be approached with
care. The two basic assumptions of the model of Jenny et al. (1949) and
Olson (1963) are that the litter accumulation has reached an equilibrium
state, where annual accession equals annual decomposition, and that the
annual litter fall is constant from year to year.
Where predicted litter half-lives are short, the assumption of
steady state, if substantiated by knowledge of fire history and seral
status, is probably valid. However in arid and montane regions, where
litter half-life may be several decades (such as Sierra Nevada pine
forests, California; Jenny et al. 1949), caution is necessary. A factor
causing large variation in litter fall from year to year is branch fall due
to high winds (Christiansen, 1975). Calculation of the decomposition
constant for leaf material only, will avoid this problem, while providing
a parameter allowing comparison of decomposition rates at different sites.
The decomposition constant and litter half-life have been cal-
culated from data provided in some Australian studies, and are shown in
Table 6.5 with values from other work, for comparison. Where leaf half-
lives could be calculated, this was done. TABLE 6.5 Predicted litter half-lives in some Australian and tropical American forests. DS : dry sclerophyll WS : wet sclerophyll RF : rainforest
Locality Dominant Forest Litter Leaf litter Reference species type half-life half-life (yr) (yr)
Dwellingup, W.A. E. marginate DS 5.1 3.3. Hatch, 1955 (mature, 22 yrs. since fire)
Mt. Lofty Ra., E. obliqua, DS 2.9 Lee & Correll, 1978 S.A. . E. baxteri
N. Stradbroke I., E. signata WS 2.9 2.0 Rogers & Westman, 1977 Qld.
Nr. Brisbane, Qld. E. umbra WS 2.3 1.0 Birk, 1979 E. baileyana (including (overstorey Angophora grass) leaves only)
New England, N.S.W. Eucalyptus spp. WS 0.25 Richards, 1976
New England, N.S.W. RF 0.50 Richards, 1976
Taree, N.S.W. E. pilularis ws 2.5 1.3 van Loon, 1969 (25 yr. regrowth)
Sydney, N.S.W. E. piperita DS 8.6 Hannon, 1956, 1958
Mt. Donna Huang, Vic. Nothofagus RF 1.7 0.92 Howard, 1973 cunninghamii
Hume Plateau, Vic. E. obliqua MS 3.6 Attiwill, 1968
Hume Plateau, Vic. E. regnans WS 2.0 1.3 Ashton, 1975 mature pole WS 2.1 1.3 Locality: Dominant Forest Litter Leaf litter Reference species type half-life half-life (yr) (yr)
Kinglake West, Eucalyptus Ds 1.9 Gill, 1964 Vic. spp.
,f Wilson's Promontory, Acmena WS 1.2 Frankenberg, 1965 Vic. smithii
North-west Pas. E. obliqua, WS 4.0 Madden et al., 1976 mature 50 yr. regrowth WS 6.5 20 yr. regrowth WS 6.6 13.yr. regrowth WS 6.4 5 yr. regrowth WS 2.0
Anglers Creek, E.Tas. E. oblique, WS 0.71 0.51 Present study 0. argophylla, A. mearnsii
Colombia Broad-leaved forest 1.1 Jenny et al., 1949 Broad-leaved RE 0.41
Panama Tropical Moist forest 0.38 Golley et al., 1975 Premontane Wet forest 0.32 Riverine forest 0.85 210
A comparison of calculated and measured litter half-lives is possible from the data provided by Golley et al. (1975) from Panama.
Their graph of loss of dry weight of litter in "decomposition plots" shows that half of the dry weight had been lost after 6 months in the
Tropical Moist forest, and after 6 12 months in Premontane Wet forest.
The corresponding calculated half-lives are 0.38 years and 0.32 years respectively.
Inspection of aerial photographs taken in 1967 and 1979 (Plates
6.2 and 6.1), and the lack of charcoal on any of the large trees in the area indicate that the Anglers Creek study area has been undisturbed for at least several decades, despite nearby forestry activity. The fact that the litter layer beneath the litter bins had almost disappeared in a year indicates that rates of decomposition and litter fall were similar and that, according to the exponential model, the steady state would be reached in just over a year. Thus a forest floor equilibrium may be assumed, subject to annual fluctuations of litter fall (Witkamp and van der Drift, 1961; van der Drift, 1963). Assuming the Anglers Creek litter fall value is close to the long-term average, the estimates of litter half-life may be regarded as valid.
Comparison with calculated litter half-lives from other studies
(Table 6.5) shows that the predicted rate of decomposition at Anglers
Creek is high compared with most eucalypt forests, and in the range found in tropical forests, where extremely high decomposition rates have been found (Nye, 1961; Madge, 1965). A number of factors contribute to this situation, not the least being the composition of the litter. Firstly, eucalypt wood and bark are resistant to decomposition (Ashton, 1975); how- ever, as E. obliqua is a non-decorticating species, large strips do not drop continuously or in such large quantities as in other eucalypt species.
The two E. obliqua trees near the study area were not close enough to contribute much wood to the forest floor studied. Olearia wood and bark appears less resistant to decomposition. Secondly, Meentemeyer (1978) 211 has found that an index related to water availability, Actual Evapo- transpiration, is closely correlated with litter decomposition rate, and the even distribution of rainfall throughout the year (Figure 5.2) must result in a high value of this index at Anglers Creek. Thirdly, and perhaps dependent on these factors, a high biomass of invertebrates is present (Chapter 5 ), and this has been shown in several studies
(e.g. Perel et a/., 1971) to increase decomposition rates. The impact of the high land amphipod density on the decomposition of litter is the subject of a further study, not reported here.
Ashton (1975) found 0. argophylla richer in nitrogen, potassium, calcium and phosphorus than the dominant E. regnans in a Victorian wet sclerophyll forest, although not as rich as other understorey species,
Acacia dealbata and Cassinia aculeata. He also showed that some elements, notably nitrogen and phosphorus, are withdrawn from the leaves prior to leaf fall in both E. regnans and understorey species. Attiwill (1968) has demonstrated that a large proportion of the phosphorus is withdrawn from E. obligua leaves before they fall, in marked contrast to the behaviour of this element in hardwoods of other parts of the world. Australian soils are generally low in phosphorus (Wild, 1958; Stewart, 1959) and Specht and Groves (1966) have postulated that the removal of phosphorus from the leaves of plants in Australian heath communities before leaf-fall is an important adaptation to a phosphorus-poor environment. The high concen- trations of this element in E. obligua and Acacia litter from Anglers Creek than in similar material from the Hume Plateau may indicate its presence in generally higher levels in the environment. In another Tasmanian study,
Harwood and Jackson (1975) found concentrations of phosphorus (and of potassium and calcium) in wet sclerophyll forest floor litter, similar to those at Anglers Creek.
Direct comparison of soil phosphorus content with other Australian forests is difficult as this is generally quoted in terms of phosphate 212
only (e.g. Wild, 1958; Coaldrake and Haydock, 1958; Stewart, 1959).
Phosphorus content of Australian wet sclerophyll forest soil is higher
than that of dry sclerophyll forest (phosphate, Beadle, 1954; phosphorus,
Ellis, 1971); certainly the eastern fringe of the continent and Tasmania,
where most of the wet forest type occurs, have generally higher phosphorus
levels than the central and northern regions (Wild, 1958). Ellis (1971), who measured total phosphorus, found that the concentration of this element
in wet sclerophyll forest soil was high on the surface and decreased markedly with depth, in contrast to that of dry sclerophyll forest, which was uni-
formly low. This is most likely due to the high organic content of the
upper soil layers of the former forest type. Even so, phosphorus con-
centration in the top 5 cm of soil at Anglers Creek is eight times higher
than in the surface soil at the more phosphorus-rich of Ellis' (1971) wet
sclerophyll sites, where E. obliqua is also the dominant eucalypt.
All this evidence points to unusually high levels of phosphorus
available to plants and animals at Anglers Creek, and it is therefore
unlikely that this element is limiting here, as it often is in other
Australian plant communities, such as heath (Specht and Groves, 1966). 213
CHAPTER SEVEN
NICHE SEPARATION
A study of the ecology of two sympatric species of terrestrial amphipod
Section 7.1 Introduction
The diversity of the land amphipod fauna of Tasmania has been shown to be quite high. Many samples collected from various parts of the island contained more than one species and it soon became apparent that it was usual to find a number of species coexisting in forest floor and other habitats. In fact, 322 of a total 502 collections contained two or more species. At six spots sampled, five landhopper species were represented in the litter or moss, although the search area in each case was quite small. Two of these places were on De Witt Island, off the south coast; the others were on the Kermandie track to the Hartz Mountains, near Tahune Bridge on the Huon River, in the upper Olga River valley and on Philips Island in Macquarie Harbour. In all, thirteen species were represented in these six samples. The coexistence of land amphipod species is therefore very common in Tasmania.
Field and laboratory observations, as well as the limited literature on the subject (Lawrence, 1953; Clark, 1954; Hurley, 1968; Duncan, 1969) support the belief that the primary source of energy of landhoppers is decomposing litter. The apparent coexistence of these species over a long period contradicts the principle of competitive exclusion (Gause, 1934;'
Slobodkin et al., 1967). As a corollary to this principle, one would expect some difference in resource utilization between coexisting species, and an attempt to determine this difference between two coexisting land- hopper species is reported in the following chapter.
The closely related species Keratroides vulgaris (Friend) and
K. angulosus (Friend) were chosen for this investigation for the following reasons. Firstly, they are fairly easily distinguishable alive, and with the naked eye. Secondly, one occasionally finds extremely high densities 214 of these species in localities where both occur together (over 2 10 000/m of these two species were found on one sampling occasion), which reduces the effort required to collect statistically reliable data. Thirdly, these species are sympatric through most of the range of K. angulosus (see Figures 4.16 and 4.20).
Morphological and other differences between K. vulgaris and K. angulosus
Friend (1979) outlined the morphological distinctions between these two amphipod species. The colour difference is the most useful for this study. While K. vulgaris is a normally pigmented landhopper (although somewhat variable in colour), K. angulosus is almost completely lacking in pigment, apart from the ommatidia of the eye (Plate 7.1). While this distinction is very obvious in adults, very small individuals of all normally pigmented species are usually light-coloured, and K. vulgaris is no exception. The ultimate criterion used was the colour of the large second antenna, which individuals of both species extend ahead of themselves when walking. Those of K. angulosus are off-white, like the body, while even in a very pale K. vulgaris, the antennae retain a pink tinge.
Several other obvious differences are of interest in this inves- tigation. K. angulosus is a rather smaller species than K. vulgaris, maturing at a smaller size, the larger individuals (females) reaching a maximum size of only 8-9 mm at Anglers Creek while some K. vulgaris females 15 rom long have been found there. K. angulosus young are between 2.5 mra and 3.5 mm in length on hatching, whereas 3.0-4.0 mm is the range for the larger species. Relatively shorter peraeopods, antennae and smaller eyes also distinguish K. angulosus from K. vulgaris
(Figures 3.60: and 3.47, Plate 7.1).
The close similarity of the mouthparts of the two species (Figures
3.49 and 3.62) suggests that there is no fundamental difference in feeding habits. This is in contrast to another species of the genus, K. albus n.sp., Plate 7.1 Live K. angulosus (upper) and K. vulgaris individuals.
• • • • • • • • • • • • • • • • • • • • • MATUSHITA COMMUNICATION INDUSTRIAL CO
Plate 7.2 Pen recorder trace of activity recorder output [Section 7.2(b)]. 215 which is rarely found in litter, being a burrower, and possesses maxi-
llipeds and mandibles (Figures 3.45, 46) strikingly different from those of all other Tasmanian species. With respect to feeding differences,
laboratory observations have shown that both K. vulgaris and K. angulosus feed on decaying leaves, and apparently prefer these to other forest litter.
Casual observations made during collecting and handling of the two species in question have revealed behavioural differences which appear significant. When disturbed under the forest litter, or placed on a pile of soil in the laboratory, K. vulgaris flicks vigorously away in search of cover in a manner characteristic of many landhoppers. It repeatedly flexes and straightens its abdomen rapidly and this causes quick movement across the substrate. Unlike a sandhopper, the landhopper remains on its side while escaping, rarely regaining its upright posture between move- ments. This locomotion is closer to that displayed by many aquatic species
(marine and freshwater) when stranded, than to that displayed by the supra- littoral sandhoppers. K. angulosus individuals, on the other hand, attempt to burrow when disturbed, or if this is impossible, their flicking movement is much less vigorous than that of K. vulgaris.
An observation made during initial collecting in the general area of Anglers Creek was that light-coloured amphipods (K. angulosus) seemed to be the only ones found at deeper levels below the litter. The invest- igation of niche differences between the species began at this point, with a study of vertical separation.
Section 7.2 Methods and Results
a) Vertical separation
Two different approaches were used to investigate this aspect of habitat partitioning. The first was to determine the position in the litter/soil profile occupied by individuals of both species during the day by a gelatin-embedding method. Sedondly, pitfall trapping at different levels in the profile was carried out to see whether any habitat differences 216 were reflected in differences in the species' activity at each level over an extended period.
i) Gelatin embedding
An open-ended sheet-metal cylinder, 20 cm in diameter, was placed on the litter surface, which was cut with a sharp knife enabling the cylinder to be pushed cleanly through the litter into the soil to a depth of 10 am. The soil and litter in the cylinder was then frozen in situ by packing dry ice onto the surface of the litter inside the cylinder and in a trench dug around outside it. The cores were then removed and kept frozen in transport to the laboratory, where they were stored at 0 0 -10 C. After each core was heated to 40 C to allow full penetration of the embedding agent, a 20% gelatin solution (with thymol added as an o antibiotic) at 50 C was poured over the core, still in the sampling cylinder, sealed at the base to prevent the escape of the solution. Setting of the gelatin was carried out at 5 °C.
When the gelatin had set, each core was removed from the cylinder and divided into litter layer and 5 mm or 10 mm soil layers, to a depth of 50 mm. The separated layers were broken up in warm water and sorted through for amphipods, which were sufficiently well preserved for identification and measurement under a stereomicroscope.
The numbers of each species, found at various levels on the two sampling dates (6 May 1976 and 22 February 1977) are presented in
Figure 7.1. The data from 30 January 1976 presented by Friend and
Richardson (1977) do, not appear here because the direct method using hot gelatin solution in the field seems to drive the amphipods downward before killing them, creating false patterns of distribution. The vertical distribution on both dates shows a higher proportion of K. angu1osus individuals at lower soil levels. Table 7.1 shows that the percentage of individuals of each species found more than 1 am below the soil/litter interface is much greater in K. angulosus on both occasions.
217
TABLE 7.1 Numbers of K. vulgaris and K. angulosus found more
than 1 cm below soil surface in gelatin cores from
Anglers Creek.
Date 6 May 1976 22 February 1977
Species K. vulgaris K. angulosus K. vulgaris K. angulosus
No. below 1 cm depth 1 8 10 44
Total no, in core 9 22 104 136
% below 1 cm depth 11% 36% 10% 32%
The mean depth of individuals of each species on each date was
calculated by designating each layer with a depth value (i.e. the litter
layer, with - 0.5 cm, 0-1 cm soil layer with 0.5 cm, 1-2 cm soil layer
with 1.5 am etc.). Table 7.2 shows the mean depth at which individuals
were found, obtained in this way, and the significance of the differences
between them, compared by t-tests. The mean depths of K. vulgaris and
K. angulosus are the only significantly different ones, indicating a
vertical separation between the species. The small number of amphipods
on 6 May makes conclusions difficult : however the lack of amphipods in
the litter layer may be due to the unseasonal dryness of the previous
weeks . (Figure 5.3) compared with the weeks preceding the second sampling
date.
TABLE 7.2. Mean depth of K. vulgaris and K. angulosus individuals (cm below soil surface, as explained in text) in gelatin core taken on each date. Significance of difference of means by t-test. _ 6 May 1976 22 February 1977
K. vulgaris 0.61 n.s. 0.32 ***
K. angulosus 0.86 n.s. 0.75
Amphipods taken on 22 February 1977 were placed into 1 mm size
classes. By assigning the class midpoint length to each specimen, a K.angulosus K. vulgaris
0
1
2- 6-5 - 76 2 animals
litter layer
2-
4- 22-2 - 77 10 animals
Figure 7.1 Vertical distribution of Keratroides angulosus and K. vulgaris populations at Anglers Creek site in May 1976 and February 1977, assessed by a gelatin-embedding technique.
218
mean length was calculated for each species and each soil layer, and the
differences between the means tested using t-tests (Table 7.3). The
results indicate that although K. vulgaris individuals were larger than
K. angulosus above 1 cm depth, animals of both species found below that
level were of the same average size, and significantly smaller than those
above 1 cm in the soil. The implication here is that the size differences
between the two species is contributing towards the difference in their
vertical distribution.
. TABLE 7.3.. Means of individual lengths (mm) of amphipods at different levels in gelatin core on 22 February 1977 (levels denoted as depth below soil surface in cm ). Number at each level shown in Figure 7.1.
Level K. vulgaris K. angulosus
Litter layer 7.1 * * 5.3 I * * * n.s. I 0-1 5.2 * * * 4.1
n s.
4.6 n.s. 4.31
n s.
2-3 4.7I
n.s.I ' I 3-4 3.3
Pitfall trapping at different levels
Fifteen wide-mouthed jars containing a saturated solution of
picric acid were sunk into the ground to three different depths. A plastic
funnel (slope 600 to the horizontal), with the spout removed, rested in
the mouth of each jar so that the rim of the funnel (11 am diameter) pressed
against the wall of the hole dug for the trap. These rims were at depths
of either 2 am (in the litter layer), 4 cm (below the fermentation layer)
or 7 am below the surface of the litter. Groups of three traps (one at 219 each depth) were placed at five locations (marked A, Figure 7.2) around the study site to encompass variation within the area. No adjacent traps were closer than one metre to each other. Small covers were erected 5 cm above the openings of the traps to prevent dilution of the preservative by rainfall. Trapped animals were collected after 14 days
(on 6 May 1976).
A total of 405 K. vulgaris and 244 K. angulosus (as well as 187
Mysticotalitrus spp.) were caught over the 14-day period. Table 7.4 shows the mean numbers in traps at each level. Comparing the catches of the Keratroides species at different levels, a 2 x 3 contingency chi-squared test shows that the distribution of the two species between 2 the levels are significantly different (d.f. = 2, x = 6.59 * ). Within- species comparisons of catches at different depths revealed that the only significantly different mean catches were those of K. angulosus at
7 cm and those at 2 cm and 4 cm combined (d.f. = 13, t = 2.48 ). This indicates that while K. angulosus ranges throughout the layers sampled by the traps, K. vulgaris moves about only in the upper layers.
TABLE 7.4 Mean catches (average of 5) of K. vulgaris and K. angulosus
in pitfall traps set at different levels.
Level 1: lip of trap 2 cm below litter surface
Level 2: lip of trap 4 cm below litter surface
Level 3: lip of trap 7 cm below litter surface.
Level K. vulgaris K. angulosus
1 23.6+6.20 13.2+2.58
2 26.4±3.22 12.2±3.07
3 31.0±7.85 23.4±4.91
Another interesting finding was the significantly higher number of K. vulgaris than K. angulosus in the total catch (d. f. = 28, t = 2.62 ). The population sample taken on 28 April 1976 (during the
Modified areas A Pitfall traps • •A Population sampling Log area B x
A
10m Log • A • x Trays
• Traps
Figure 7.2 Relative location of population sampling area, litter trays and pitfall traps set at different levels [Section 7.2(a)(ii)]. Area PQ shown in detail in Figure 7.6. 220
trapping session) contained about four times as many K. angulosus as
K. vulgaris, and at no time during the 2-year study was K. vulgaris
the more numerous species (Figure 4.2). The proportion of K. vulgaris
in all Keratroides caught in 7 cm-deep traps (0.570) was compared with
this proportion in the population at large on 28 February 1976 (0.208)
by the t-test for comparing proportions (Sokal and Rohlf, 1969), and was *** found to be significantly different (d.f. = 03, t = 9.04 ). A possible
explanation is that K. vulgaris is more active in the field than K. angulosus
and this possibility was therefore investigated.
Activity differences
The possibility of inherent differences in levels of locomotory
activity displayed by the two species was investigated by monitoring
activity in the laboratory and by further pitfall trapping programmes.
Pitfall catch is a function of the activity and density of the population
at risk (Mitchell, 1963; Greenslade, 1964). In conjunction with good
population data from other sources it is thus feasible to gain estimates
of relative levels of activity.
i) Activity monitoring in the laboratory
A capacitance-type activity recorder as described by Grobbelaar
et al. (1967), using the sensor designed by Evans (1972), was employed to measure the level of activity of individual amphipods. The sensor/animal
chamber consisted of a pair of printed circuit boards forming the plates
of the condenser, held apart by a sheet of 4 mm Perspex. The space for
the animal was formed by a circular hole (10 cm diameter) in the Perspex
sheet. Relative humidity in the chamber was kept at 100% by a piece of
filter paper soaked with water, laid on the lower condenser plate, with an
Olearia leaf to provide food. The animal chamber was placed in a small
Faraday cage (to reduce electrical interference) which in turn was placed
in a cabinet kept at a constant 100C. For most runs (see Table 7.5) a
light in the cabinet connected to a time switch provided 10h light and 14h
221
darkness, corresponding to day length in the field at that time.
TABLE 7.5 Hourly activity of individual K. vulgaris and K. angulosus
measured in the laboratory. Activity is expressed in
arbitrary counts, being the product of duration and intensity
of activity in each hour, averaged over the duration of the
experiment. "Light/dark" indicates 10h light, 14h darkness.
Species Weight (mg) Activity units/h Conditions Duration (h)
K. angulosus 10.9 7.12 light/dark 34
9.0 5.08 light/dark 24
7.0 2.15 dark 33
K. vulgaris 8.3 11.89 light/dark 28
4.6 14.29 light/dark 21
Amphipods were placed singly into the chamber soon after
collection from Anglers Creek. After the experimental run (not longer
than 351) each animal was weighed alive on a Sartorius '2400 analytical
balance.
The output from the detector caused deflection of a National
VP 654B pen recorder, and a sample trace is shown in Plate 7.2. The
trace was analysed by ascribing a value on an arbitrary 0-5 scale separately
to the amplitude and duration of the activity traces during each hour.
The product of amplitude and duration scores each hour gave a measure of
overall activity. Figure 7.3 shows the variation in activity of two
amphipods over the experimental period. The mean activity per hour of
each of these animals is shown in Table 7.5, and it is obvious that
the K. vulgaris shows a higher level of activity than the K. angulosus,
despite their similarity in weight. The results of further experiments
(Table 7.5 indicate that K. vulgaris is the more active species. The
activity record also shows a increase in activity of both species during
dark periods (Figure 7.3). Unfortunately, after these runs, the activity
25- K. vulgaris
15-
0 -6- c 0
25 - K. angulosus 4-
.- 4- < 15-
1 &CO 18-0() Time of day (h)
Figure 7.3 Hourly activity of a K. vulgaris individual (weight 8.3 mg) and a K. angulosus individual (weight 9.0 mg) under 10h: 14h light/dark cycle, at 10°C. Activity is expressed as the product of the amplitude and duration of activity during each hour, each assessed on an arbitrary 0-5 scale. Arrows indicate the interval of measurement, while the solid (dark hours) and open (light hours) rectangles describe the artificial photoperiod regime. 222 recorder stopped working satisfactorily and no further data were collected.
ii) Day-night pitfall trapping
Another trapping programme was carried out to investigate the daily patterns and relative levels of surface activity of K. vulgaris and
K. angulosus in the field. Two sets of data were collected at Anglers
Creek, one in August 1978 and one in February 1979, in an attempt to assess any seasonal differences.
Twelve pitfall traps were placed in the population sampling area Hon 1 August 1978, a week before the initial trapping period, to reduce the "digging-in effect" (Greenslade, 1973). The traps consisted of one plastic drinking cup inside another, set into the ground so that the inner cup, containing saturated aqueous picric acid solution, could be removed and emptied frequently without disturbing the walls of the pit.
The rim of the outer (250 mL) cup was trimmed and the cup set into the forest floor so that the lip was just below the soil surface. Holes in the base of this cup allowed the escape of rainwater when the traps were closed. The inner cup, of 225 mL capacity, fitted into the larger one with its rim (7 cm diameter) flush with the soil surface (Figure 7.4).
Inner cups with perforated bases were filled with soil and inserted into
the outer cups to close the traps between trapping periods. Rainfall shelters were not used because of the short trapping intervals. After collection, trapped animals were transferred to 70% alcohol (with glycerol) for preservation.
In August, traps were opened at dusk on the 5th and emptied at dawn, dusk and midnight the next day, then at dawn, noon and dusk on the
7th. In February, traps were opened at dusk on the 23rd, emptied at dawn and dusk on the 24th and at dawn on the 25th. A Casella thermohygrograph provided a record of temperature and humidity during both trapping sessions.
Table 7.6 summarises the results of this trapping programme.
During the day, fewer individuals of each species fell into traps each Figure 7.4 Design of pitfall trap used in day-night trapping, to allow repeated emptying and resetting. Inner plastic cup may be removed from outer cup and replaced with minimal disturbance to adjacent soil and litter.
K. angulosus K. vulgaris
14:
12:
10-
E - -- 8- -c 4- - Er 6- w . _J 4-
2-
5 animals
Figure 7.5 Size-frequency diagrams of K. angulosus and K. vulgaris caught in day-night pitfall programme in August 1978. 223 hour than at night, (Table 7.7) when these catches were compared by the method outlined below. This finding supports indications of low daytime activity of K. vulgaris and K. angulosus in the laboratory. There was no evidence of different catch rates before and after midnight, or before
and after noon in August when these were measured.
Hourly catch declined between the first and second nights in
the February trapping period, evidently as a result of trapping out a
large number of active individuals and thus reducing their density in the
immediate vicinity of each trap. A similar effect was noticeable, but
less narked in August. This obscured the differences between night and
day catches of each species. However, if we assume that over the trapping period, the catch declines linearly with time (which is only an approxim-
ation if fewer animals are removed during daylight hours) then night and
day catches can be compared by calculating the ratio of hourly catch on
the first and second nights pooled, to the hourly catch on the first day.
Approximating, if X is the catch depletion over 12 h, then the average
hourly catch for the first and second nights, n, and the hourly catch for
the first day, d are
N N-2X n = 1/2 T2- + 12 where N = the first night's catch N-X = 12 D-X and d = 2 where D = the first day's catch, had there been no previous trapping
respectively. Now the ratio of the night's to the day's hourly catch
N -X which thus compares D-X nightly and daily catches on the same terms (i.e. as if both were
caught in the second 12-hour period).
Values for i for each species over the two trapping sessions
are shown in Table 7.7. Hourly night catches exceeded hourly day catches
for each species in summer and winter, after accounting for the effect of 224
TABLE 7.6 Catches in day/night pitfall trapping programmes
in August 1978 and February 1979, showing duration
of day and night trapping periods.
5-7 August 1978 R.H. 100% Temperature 5-9 °C
1st night 1st day 2nd night 2nd day
K. vulgaris 9 4 7 2
K. angulosus 11 2 10 2
Mysticotalitrus 9 1 5 1
Duration (h) 13 11 12 11
23-25 February 1979 R.H. 95-100% Temperature 10-15 °C
1st night 1st day 2nd night
K. vulgaris 84 31 56
K. angulosus 11 6 4
Mysticotalitrus 35 25 24
Duration (h) 9.5 13.5 10.5
TABLE 7.7 Ratio of numbers caught each hour in the dark hours
to numbers caught in the light hours; see text for method
of calculation.
August 1978 February 1979
K. vulgaris 1.8 3.0
K. angulosus 4.7 1.7
Mysticotalitrus 6.2 1.6
depletion.
Therraoh_ygrograph records showed than in August, relative humidity remained at 100% for the entire trapping session, and in February dropped below this value for only a short time, reaching a minimum of 94% at 1215 on the 24th. Rainfall records from Toams Lake show that while 225
32 mm of rain fell over the 3-day trapping period (up to 0900 on
8 August), only 1 mrn was recorded during the two days of trapping in
February. The high humidity over the second period was due to low cloud which enshrouded the area in mist much of the time.
McColl (1975) found that pitfall catches of some macroarthropod groups in Nothofagus forest in New Zealand increased under conditions of high humidity over 14-day periods. The surface activity of humidity- sensitive animals like amphipods might be expected to decrease in low humidities. It is interesting, therefore, that diurnal activity cycles are apparently found under conditions of almost constant high humidity.
This indicates that, as in the laboratory, landhopper activity in the field is increased in low light intensities. Summer catches of K. vulgaris *** ** (d.f. = 11, t = -4.75 ) and Mysticotalitrus (d.f. = 11, t = -3.40 ) were compared with the respective winter catches by a paired sample t-test, and found to be significantly higher. K. angulosus catches, however, remained at a low level in both seasons. Annual changes of population
density (Figure 8.2) are not sufficiently large to explain this phenomenon;
the most probable explanation is that the activity of the litter-dwelling
groups (K. vulgaris and Mysticotalitrus) is increased with temperature, but that in K. angulosus, this increase either does not occur, or, more
likely, is not expressed as increased surface activity.
The amphipods caught in August 1978 were measured, and Figure 7.5
shows the structure of the trapped populations of K. vulgaris and
K. angulosus. When compared to the structures of August populations
determined by random sampling (Figure 8.1 ), the distribution of trapped
animals of both species shows that animals caught in pitfalls tend to be
larger, on average, than those in the population at large. By assigning
the size-class midpoint length to each trapped animal and to each animal
taken in August 1977 random samples, the mean length of animals of each
species taken in these two ways could be compared. In both K. vulgaris *** *** (I.f. = 85, t = 4.64 ) and K. angulosus (d.f. = 197, t = 6.44 ) the 226 lengths of trapped and random-sampled animals were significantly different.
c) Colonization tendencies
One advantage held by a more active species over a less active species in a guild is the ability to move first into newly-available areas. In an ecosystem where new habitat space is constantly becoming available, r-selected opportunistic species can survive by rapid colon- ization, despite being replaced later by more specialist K-selected species (MacArthur and Wilson, 1967). In other situations, the first
colonist is at a distinct advantage, which may allow its persistence
even after the arrival of-better adapted species (Sale, 1977).
In view of the apparently greater activity of K. vulgaris than
K. angulosus, several experiments were carried out in an attempt to measure the relative speed at which these species appeared in unpopulated areas. These experiments may be divided into two categories: those in which pitfall trapping was used to assess amphipod activity in the areas concerned, and those in which soil and litter samples were removed to measure the presence of the two species in newly-available areas.
i) Pitfall trapping in scalded and cleared areas
Interpretation of pitfall trapping data is subject to a number of difficulties, not the least of which is variation due to size and weather conditions (Greenslade, 1964; Duncan, 1969). In the design of a trapping programme to investigate differences in activity of K. vulgaris and K. angulosus in the centre of modified areas of forest floor, an attempt was made to account for this variation. The total area available for this exercise was limited, as area B (Figure 7.6) was still reserved for population sampling, and outside it, clear areas of forest floor were less common. On the other hand, for the purposes of extrapolating population sampling data to pitfall areas it was desirable to trap fairly close to the area to which it applied. The arrangement of traps used is shown in •C4
Population sampling area
Slope j
.C23 VIS2 C2A •
Figure 7.6 Location of control pitfall traps and traps in areas scalded or cleared of litter, at Anglers Creek site. Reference codes are explained in Section 7.2(c)(i). 227
Figure 7.6 in which the line PQ runs along an area open under Olearia and Eicksonia, which appears very similar to the area B.
The traps were of the same design as those used to compare activity at different levels [section a)ii)]. The plastic funnels were positioned at 4 cm below the litter surface (in the fermentation layer) and so catches were influenced mainly by activity in the litter and on its surface. The pitfall traps were set up as follows:
- Five control traps (Cl, C2A, C2B, C3 and C4) were set up near
modified areas to provide information on amphipod activity in
the litter.
- Two traps (NS1, WS2) were placed in the middle of areas in
which all amphipods had been killed by pouring 5 L of boiling
water into an open-ended sheet metal cylinder (diameter 32 cm)
which had been pushed 5 cm into the forest floor. This water
took about one minute to drain away. A test showed that no
amphipods in the 32 cm diameter area survived this treatment,
and that a number immediately outside the area also died.
- Seven other traps were set up in the centre of circular areas,
which had been completely cleared of litter, of the following
diameters: 2 in (trap X), 1 in (traps Y1 and Y2) and 40 cm
(traps Zl, Z2, Z3 and Z4). In the 2 in area, two other traps
were placed 50 am (XA) and 75 cm (XB) from the centre trap.
Trapping was carried out over four periods of fifteen days between 11 August and 23 October 1976. This programme was so designed to provide sufficient data from the small number of traps and to monitor any long-term trends.
The catches of K. vulgaris and K. angulosus during each 15-day period are shown in Table 7.8. Pitfall catches of both species in the scalded areas were initially lower than in adjacent control traps, but slowly increased during the progrcuunte. This effect might be expected as the populations in the scalded litter rose towards their previous
228 TABLE 7.8 Catches of K. vu1g4r,t4 And K. an9u1osus in pi:.tfall
traps in modified and control areas, August - October
1976. See text for full explanation. 2 Date 1 : 11.viii - 26.viii Population density (Chapter 8)/m
2 : 8.ix - 23.ix K. vulgaris K. angulosus
3 : 23.ix - 8.x 11.viii 651 3168
4 : 8.x - 23.x 8.ix 846 2300
8.x 1063 2821 Control traps
Cl C2A C2B C3 C4
Date K.v. K.a. K.v. K.a. K.v. K.a. K.v. K.a. K.v. K.a.
1 38 27 32 17
2 45 45 29 22
3 31 94 53 • 69
4 31 80 21 84 24 9 14 7
Cleared areas 40 cm diameter Zl Z2 Z3 Z4
Date K.v. K.a. K.v. K.a. K.v. K.a. K.v. K.a. 1 71 75 37 27
2 31 26 29 47 13 19 8 36
3 28 31 27 26 20 19 3 8
4 5 1 3 7
1 m diameter Y1 Y2 2 m diameter X Date K.v. k.a. K.v. K.a. Date K.v. K.a. 1 8 51 34 37 1 33 59
2 4 2 9 8 2 2 2
3 3 5 7 9 3 10 3
Scalded areas XA 2 XA 3 2
WS1 WS2 XB 2 2 2 Date K.v. K.a. K.v. K.a.
1 21 19 12 5
2 16 25 19 8
3 36 73 40 17 229 levels. Catches of both species changed in the same way; however in the initial catches of both control and scalded-area traps, the higher number of K. vulgaris than K. angulosus in contrast to their relative population numbers bears witness to the greater surface activity of the larger species.
An interesting effect appearing with respect to catches in most of the longer-standing traps with litter around them (Cl, C2A, WS1) is the steady increase in the ratio of K. angulosus to K. vulgaris. This seems to be a real effect, although it was not observed at WS2. It is possibly due to development over six to eight weeks of some condition at or near the trap which is more attractive to K. angulosus than to
K. vulgaris.
Removing litter around traps decreased catches of both species over several weeks.
ii) Recolonization of litter
In this series of experiments, the amphipods in circular plots of 32 am diameter were killed with boiling water as described in i); these plots were sampled after several days or weeks by removing cores of soil and litter, and searching through these in the laboratory for live amphipods. Two variations to this procedure were employed: the litter was removed before scalding, and the area was either left this way, or pieces of black polythene sheet cut to the approximate size and shape of
Olearia leaves were placed on the scalded plot to provide shelter but not food. Where these variations were used, and on several scalded plots with litter, a square of birdwire (mesh size 1 x 1.5 cm) of side
45 am was pegged loosely to the ground over the area to prevent litter from falling or blowing into it.
In April and Nay 1976, five plots were scalded, and later sampled, two after a week, two after a fortnight and one after four weeks.
The removal of cores 7 cm in diameter from near the centre of each plot 230 allowed the amphipod population to be assessed. The number of cores taken from each plot was increased from three to six and seven during the course of the study as low amphipod densities were found.
Between March and June 1977, nine experiments were carried out: in three the scalded litter was left intact, in three it was removed, and in a further three, it was replaced with polythene leaves as described . above. Sampling this time was by removal of one core 20 cm in diameter from the centre of each plot, after two or five weeks.
With the small number of experiments performed, each of which took a considerable amount of effort and destroyed some amphipod habitat, no clear trends due to time could be discerned, so the results of all similar experiments were pooled. Table 7.9 shows the numbers of each species found in scalded plots under each treatment, irrespective of the time for colonization and number of cores taken. The total number of each species found in all plots also appears in this table.
TABLE 7.9 Numbers of amphipods found alive in areas of forest floor
which had been treated in various ways then left for one
to five weeks before sampling.
Treatment Area Sampled No. of K. vulgaris K. angulosus Mysticotalitrus (m2) Experiments
Scalded area with litter 0.210 8 30 3 0
Scalded area with litter removed 0.094 3 9 3 0
Scalded area, litter re- placed with polythene leaves 0.094 3 17 0 3
All experi- ments 0.398 14 56 6 3
Despite its relatively low numbers in the surrounding area,
K. vulgaris appears from this pilot study to make up a very large part of the population in newly-available areas. Pitfall trapping in scalded 231 areas [section c)i)] shows that K. angulosus individuals are active in the litter within the first two weeks, and that this activity increases after four weeks. The higher numbers of K. vulgaris in re- colonized plots during the day may be an indication of wider physiological tolerance of this species.
An interesting feature is revealed by the number of K. vulgaris found under polythene leaves, which is comparable to the number under scalded litter, when the area sampled is taken into account (Table 7.9).
The shelter provided by litter and polythene leaves appears to be more important in attracting K. vulgaris individuals than their respective nutritive values.
d) Desiccation tolerance
The integument of the landhoppers is relatively permeable to water, and as the gills are external and are always kept moist, these amphipods are very susceptible to desiccation (Hurley, 1968). Hygrometer records made near the litter surface (Chapter 5) showed that in all months of the year, relative humidity dropped below 100% on most days and that on summer nights the maximum value reached was often below 90-95%.
If, as suggested by the experiments reported above, K. vulgaris and
K. angulosus display differences in vertical distribution and surface activity, one might expect different tolerances of low humidity levels such as those experienced on the litter surface. Laboratory experiments were carried out to investigate this possibility.
Solutions of sodium hydroxide at graded concentrations, or water maintained the relative humidity at chosen levels in experimental cells.
These cells were constructed of two plastic Petri dish bottoms, the lower one containing the liquid and the upper one inverted, with fine nylon mesh (25 meshes/cm) glued across the opening, forming a floor for the 232 amphipods, which were introduced through an 8 mm diameter hole drilled in the upper Petri dish. This hole was sealed during experi- ments and equilibration. The mesh was glued to the Petri dish with elastic cement (Selley's Silastic) on a glass plate so that the lower surface was flat and formed a good seal with the lower Petri dish, with the aid of paraffin grease.
The actual relative humidities produced by the sodium hydroxide solutions were measured after equilibration for 24 h with a humidity probe inserted through the hole in the upper Petri dish. The concentrations of solutions used and the measured relative humidities of air above them at 100C are shown below.
Concentration of Relative
solution (Ww/w NaOH) humidity (%)
Water 100
0.5 98
1.0 95
2.5 90
The cells were placed in darkness in a constant temperature o cabinet at 10 C. Soil and litter containing amphipods was collected from Anglers Creek and stored for up to two weeks at 5 °C. Before experi- ments, amphipods were sorted into species, then kept for 24 h at 10 °C in closed 2 L ice-cream containers with a damp plaster-of-paris floor, and with several decomposing Olearia leaves for food.
Amphipods were introduced into the cells at the beginning of each experiment, and the holes were sealed. The size structure of the animals chosen for each cell represented approximately that of the population at large. At each inspection (at various time intervals shown in Figures 7.7 and 7.8), dead animals were recorded and removed from the cells, then pre- served 6o that their identity could be checked. Torpid animals were prodded and if there was no movement in response, they were judged dead. 233
Two series of experiments provided the data on the percentage survival of the two species at various humidities which averepresented in Figures 7.7 and 7.8. In May 1976, two cells at each of the four relative humidities were set up and about 10 animals of each species placed in each one. Fifteen to thirty-one amphipods of each species were introduced into each cell in March 1978. The first experiment ran for 72 h and the second for 96 h.
The results of the two experiments in terms of both the survival and time to 50% mortality (Table 7.10) are somewhat conflicting. In the first series of tests, the survival of both species over 72 h was good at
100% R.H; however, at 98%, K. angulosus individuals died much sooner than
K. vulgaris. Similarly at 90% and 95% R.H., the survival of K. angulosus was much poorer.
Survival rate at 100% R.H. was lower in both species in the second series of tests. On the other hand, at 98% and 95% R.H., the effect of reduced humidity was less marked in both species than in the first series.
The greatest difference between the two species this time was at 90% R.H., where all K. angulosus died much more quickly than all K. vulgaris: however in both groups, 50% mortality took more than three times as long as it did in the first set of experiments. Table 7.10 also shows the numbers of animals in each ,cell in the two series. The most probable reason for better survival at lower humidities in March 1978 is that the larger numbers of animals in each cell (particularly K. angulosus) raised the humidity above that to which the solutions should have reduced the air.
At this time the humidity probe was not working satisfactorily and so the relative humidity in each cell was not measured. Amphipods were often observed huddling together, which serves to raise the R.H. in the immediate vicinity of the animals. The reason for lower survival of both species in saturated air in this experiment is unknown.
The conclusion may be drawn, however, that at least in the region 100
80
60 K. vulgaris
40
20
>
100
K.anguiosus 80
---- H 2 0 60 o 0 .59/0 NaOH
•—• 1.0% NaOH 40 2.5% NaOH
20
10 20 30 40 50 60 70 80 Time (h)
Figure 7.7 Survival of K. vulgaris and K. angulosus individuals in first desiccation tolerance experiment. Each line refers to a group of amphipods kept over either water or a sodium hydroxide solution of a particular concentration (as indicated by the legend). Relative humidities measured over these solutions are shown in the text. K. vulpHs 100
80-
60-
0-0
40-
— 20-
K. angulosus (" 100
80-
60-
40-
20-
2b 30 1+0 50 60 70 80 40 Time (h)
Figure 7.8 Survival of K. vulgaris and K. angulosus individuals in second desiccation tolerance experiment. Legend as for Figure 7.7 234 of the lowest relative humidity employed (90%) the ability of
K. vulgaris to withstand its effects is better than that of K. angulosus.
TABLE 7.10 Time to 50% mortality (h) of groups of K. vulgaris and
K. angulosus kept over NaOH solutions at various concen-
trations. Numbers of amphipods in each humidity cell
shown in brackets.
Solution First experiment Second experiment
K. vulgaris K. angulosus K. vulgaris K. angulosus
H O >72(10) >72(11) >96(24) >96(26) 2 0.5% w/wNaOH >72(10) 23(10) >96(17) >96(28)
1% w/wNaOH 16(10) 13(12) 65(24) 85(31)
2.5% w/wNaOH 14(10) 4(10) 43(24) 38(27) 235
Section 7.3 Discussion
A number of ecological, behavioural and physiological differences between K. vulgaris and K. angulosus have been elucidated. On average;
K. angulosus is found deeper in the soil and is much less active than
K. vulgaris. These differences can be related, partly at least, to the
smaller adult size of K. ablu_losus , but laboratory measurements of activity have indicated the presence of a size-independent behavioural component.
The significantly higher weight-specific oxygen consumption of K. vulgaris
(p<0.01 at 5°C, p<0.05 at 10 0C, p<0.001 at 150C; see Chapter 9 ) also points to a fundamental difference in activity. The greater ability of K. vulgaris to survive exposure to reduced humidity is in keeping with
the microhabitat difference, and with the fact that it is found more commonly and in drier places than K. angulosus (Chapter 4). The smaller size of
K. angulosus also means that, on average, it has a higher surface-to-volume ratio, thus tending to lose moisture more quickly in dry air. The more stable humidity regime of the soil (gallwork, 1976) provides a less exacting environment for this species than the litter on its surface, where humidity
is rather variable (Chapter 5).
Brood size of amphipods is usually related to the size of the individual female, within each species (landhoppers; Duncan, 1969; Saito and Kudara, 1972: supralittoral talitrids; Louis, 1977; Williams, 1978: littoral and marine species; Steele and Steele, 1969; Vlasblom, 1969;
Sheader and Chia, 1970; Van Dolah et al., 1975). While maximum brood size is smaller in K. angulosus than in K. vulgaris, females of this species carry more eggs than K. vulgaris females of the same size (Chapter 8) as their eggs are slightly smaller. As an additional compensation for smaller size, it is possible that the mortality of K. angulosus young due to desiccation is lower than that of K. vulgaris young, living as they do in a more stable humidity regime.
While both species display reduced movement over forest floor 236 cleared of all litter, K. vulgaris is much quicker to colonize scalded litter and the soil beneath polythene leaves. The preference of this species for these two altered forms of habitat seems about equal, and it may be that K. vulgaris, being very mobile, is able to select daytime habitat for shelter only, feeding elsewhere at night. These findings could indicate the presence of competition for space in the optimal litter habitats.
Morphological and behavioural differences support the hypothesis that K. angulosus occupies a more sedentary, soil-burrowing niche in contrast to the surface-roaming, litter-based existence of K. vulgaris.
Smaller size enables easier penetration into soil pore spaces, and shorter appendages are more useful in burrowing than the long slender legs and antennae of typically litter-dwelling forms. Sand-burrowing talitrids
[Talorchestia) also possess short appendages. Lack of body pigment and reduced eyes are characteristic of the animals of the soil layers (Kuhnelt,
1976), and it is interesting to note than another Keratroides species,
K. albus sp.n., which is a soil burrower (Chapter 3) also lacks body pigment and has reduced eyes and shorter, stronger appendages. Another striking similarity between K. angulosus and K. albus sp.n. is the very rounded dorsal surface of the cephalon (Figures 3.60 and 3.42). This is somewhat reminiscent of the anterior morphology of iulid millipedes especially with respect to the collum, which has been developed, according to Manton (1954) to provide the means to push through the soil. Although escaping animals burrowing in loose soil use their peraeopods to work their way down, it is tempting to speculate that the rounded head facilitates pushing through more cohesive soil and into small crevices, possibly affording more protection . for the delicate antennae than would the more gently rounded head of
K. vulgaris. K. albus sp.n. is frequently found in discrete burrows in clay. This is possibly due to the nature of the substrate rather than to different burrowing behaviour, as K. angulosus habitat is nearly always in 237 friable soil in which burrows would tend to collapse. K. albus sp.n. is also found at deeper levels in the soil than K. angulosus.
Niche differences
The problem of explaining the coexistence in the soil of a large number of species with apparently low trophic specialization has been tackled by Anderson and Healey (1972) who formed three hypotheses:
al There is an excess of food available for decomposers. Their populations are held below the carrying capacity of the energy source
(detritus) by other physical or biological factors,
b) Undetected differences exist between the species in their utilization of resources.
c) There are undetected microhabitat differences between the species.
In addition, Marcuzzi (in discussion, following Anderson, 1975b) and Wallwork (1976) have proposed that
d) Competition for food may be avoided by utilization of resources at different times. a) Regulation below food limits
Hairston et a/. (1960) have pointed out that decomposers must be fully utilizing the annual litter fall in systems where there is no net accumulation of detritus. This may be true, but it does not necessarily follow •that the detritivore fauna is fully exploiting the food available to it.
In systems where litter is not accumulating, microflora and physical effects produce breakdown not accounted for by the fauna. While several estimates imply full utilization of litter by detritivores, many suggest that this consumption is far below the annual input (Crossley,
1977). Dudich et al. (1952), for instance, concluded that the total annual litter fall passed through the digestive systems of the soil animals, 238 on the basis of a survey of 23 forests: Similarly, Nef (1957) calculated that the mesofauna alone (presumably in European temperate deciduous forests) could comminute the entire annual litter fall, and Satchell and
Lowe (1967) predicted a similar consumption by the earthworm population in a temperate deciduous woodland.
However, Bornebusch (1930) while underestimating the density of collembola and mites (Nef, 1957) calculated that the cryptozoa consumed
20-25% of the annual litter fall in a range of Danish woodlands. UlriclOs estimate (1933) of 28% was of a similar order. In an experiment using napthalene to exclude animals from litter bags, Crossley and Witkamp (1964) found a minimum loss of litter dry weight of 15% (Gist and Crossley, 1975) due to the presence of cryptozoa, while a total weight loss of 60% occurred during the first year of decay. This result was confirmed by Witkamp and
Crossley (1966) in another experiment.
Using a computer model based on biomass data from a southern
Appalachian deciduous forest and feeding rates from radiotracer studies
(Xowal and Crossley, 1971), Gist and Crossley (1975) estimated the consumpt- ion of calcium by the detritivores as 20% of the annual input to the forest floor. Consumption of potassium (7.5%) was much lower, probably because of early loss of this highly leachable element from the litter. In exclusion experiments conducted in Australian forests, Wood (1971) found that soil and litter animals caused between 6% and 31% of the annual decomp- osition of eucalypt leaves, while Macauley (1975) found the effect of exclud- ing the cryptozoa on leaf decomposition rate to be insignificant.
These low consumption figures may give the impression that all the remaining litter is potential food for detritivores. This is not true.
Substantial early losses of leachates from litter have been recorded and in fact this appears necessary before attack by animals will occur (Edwards et al., 1970). However, if these lower estimates are correct, it is most likely that a considerable amount of litter is broken down by fungi and physical processes which would have been consumed had higher cryptozoan 239 populations been present.
Thus it appears that in many forests, an excess of food for detritivores may be present. It is probable that short periods and localised areas of limited food supply occur (Anderson and Healey, 1972) especially in forests with a highly periodic litter fall, but these are probably limited in duration and extent, and could be withstood by migration of some groups, or by the temporary cessation of feeding mentioned by
Anderson and Healey (.1972).
Australian studies of the proportion of the annual litter fall con- sumed by amphipods have shown that there is an excess of food present, in the Absence of other detritivores with a similar impact on the litter fall.
Estimates, in energy terms, of 2.9% annual litter fall consumption by
Mysticotalitrus tasmaniae (Friend, 1975), 25% for Arcitalitrus sylvaticus
(Clark, 1954) and >20% for Keratroides vulgaris and K. angulosus (unpub..data) indicate such an excess. On the other hand, Duncan's study (1969) of two
Orchestia species in cocksfoot grassland in New Zealand revealed that dis- appearance of most of the litter by autumn coincided with a sharp population decline. At Anglers Creek, the only time when food might be in short supply is in early summer, if all the leaf litter on the forest floor is recently fallen and not yet sufficiently leached. This is unlikely, however, as the minimum standing crop measurement (Chapter 6) was almost half the maximum.
Under these conditions of excess food, populations must be regulated below the level which the food supply could support. Coexistence of several species of larval Tipulidae (Freeman, 1967) and of stem-boring insect larvae (Rathcke, 1976) in the presence of excess food has been shown, although no other means of population regulation were detected. Anderson
(1975b) suggested that predation, competition for oviposition sites, or climatic factors may keep the detritivore populations below the levels where competition for food might occur. Sheals (1955) and Dindal et al.
(1975) removed some predators from soil mesofauna communities by use of 240 insecticide, and found population increases in some prey species.
Regulation of prey by predators in forest litter was also demonstrated by Clarke and Grant (1968) who, by physically removing spiders from forest litter, caused a marked increase in millipede and collenbolan densities.
Parasitism may also hold populations below the carrying capacity of their environment. When parasitism of psyllid nymphs was reduced due to abnormally cold weather, the population of these bugs increased dramatically until food and oviposition sites became limiting (Clark, 1964).
Duncan (1969) found evidence of some mortality in landhoppers due to desiccation and, more commonly, to osmotic stress from submergence in rainwater. He found predation a more important cause of death in moulting animals than in others.
At Anglers Creek, the most likely controls on amphipod numbers are weather, especially very dry and very wet periods, and the limited amount of habitat space. These two controls probably interact, since animals forced into the upper litter by competition for space are more susceptible to effects of weather. In periods of low relative humidity, there is the danger of desiccation as well as the reduction of sufficiently moist leaf litter to supply food for the whole population. In very wet periods, amphipods may be caught by surface tension effects in drops of water trapped between leaves. Predators, although taking some portion of the population production, appear to be too few in number to exert any regulatory effect.
No parasitism or disease (c.f. Duncan, 1969) has been detected in either of the populations.
Consequent to the findings and discussions Above, three hypotheses are proposed.
1) Food availability is not important in controlling amphipod numbers.
2) Amphipod numbers are controlled by the available habitat space.
3) Predation is not important in regulating amphipod numbers. 241
Several experiments to test these hypotheses are outlined below.
The response of amphipod population numbers to the following habitat manipulations should be monitored over at least one year.
Experiment 1 : The addition of food without substantially increasing habitat space could be accomplished by adding moderately finely-ground decomposing litter to the litter layer.
Experiment 2 : Increase of habitat space without increasing food availability could be achieved by the addition of artificial litter to the litter layer and by increasing the depth of the friable upper soil layers.
These manipulations are crude and may not produce the desired changes in resource availability. While increases in amphipod numbers would strongly indicate a relationship between the availability of the resource and population numbers, decreases or no change could be due merely to the failure of the experimental method to provide the desired conditions.
Experiment 3 : Investigations in enclosures, either by excluding vertebrate predators with wire mesh, or by physically removing invertebrate predators following the methods of Clarke and Grant (1969), or both, should provide information on the role of predation in the regulation of amphipod numbers. b) Differences in food utilization
Although it has been known for some time that there are major trophic groupings amongst soil animals, the diversity within these groups is so large that competition between species seems inevitable (Wallwork,
1976). However, as more information on the ecology of soil animals becomes available, it appears that there are subtle differences in feeding patterns, even between detritivore species. Anderson and Healey (1972) detected some difference in gut contents between representatives of two genera of large Collembola, while Anderson (1975a) found that the diets of adults of several Cryptostigmata species could be separated to some extent. 242
McMillan (1975) compared the gut contents of three small Onychiurus
species (aollembola) and found selectivity in their feeding, as well
as small but significant differences between the species. Differences
in palatability of leaves of various plant species between four isopod
and four diplopod species were found in tests by Neuhauser and Hartenstein
(1978).
In a study of the diets of several lake-dwelling triclad species,
Reynoldson and Davies (1970) found that considerable differentiation in
diet was necessary for coexistence. Although the same categories of
food items were present in the diets of all species, the triclads showed
specialization to the extent that one category of food was taken by each
species at least 30% more often than by any other species. The differences
in diet listed above, and those found in coexisting species of woodland
pulmonates by Mason (1970) and by Jennings and Barkham (1975) involved
much greater overlap than this at the level of discrimination used by the
investigators. It has been pointed out by Pianka (1978), however, that
significant overlap on two niche dimensions may still allow niche separation.
Jennings and Barkham (1975) suggest that feeding zone may be the other
dimension separating the niches of the woodland slug species investigated.
Selective assimilation of food components may avoid competition in
some groups. Differences in enzyme complement in cryptozoans, or in their
.gut flora, have been found amongst oribatids (Luxton, 1972) and collembola
(von Tome, 1967a, b; Zinkler, 1971), litter-dwelling dipteran larvae,
diplopods and oligochaetes (Nielsen, 1962) while differences in peroxidase
activity amongst groups of species of oligochaete, diplopod, isopod and
pulmonate were demonstrated by Neuhauser and Hartenstein (1978). While
enzyme differences between species selecting different dietary components
might be expected, the possibility of these differences allowing a re-use
of food by interspecific coprophagy, as suggested by Anderson and Healey
(1972) cannot be discounted. Little field work has been done to determine
the extent of this mode of feeding. However, Nicholson et a/. (1966) 243 found the faeces of smaller animals in place of decomposed Glomeris marginata (Diplopoda) pellets and noted the presence of mites, Collembola, isopods and enchytraeids on faecal samples. Furthermore, populations of animals active in the soil water of subtropical rainforest in New South
Wales were maintained on faecal pellets of the amphipod Arcitalitrus sylvaticus by Clark (1954). While ostracods and aelosomatids apparently grazed macro-organisms from the surface of the pellets, copepods, enchytraeids and naidids ingested small particles of faeces. Coprophagy is thus another means of partitioning the detrital food source and reducing competition for
food when and where it is limited.
There is no direct evidence of feeding differences between the
Keratroides species and although this aspect was not investigated thoroughly, preference tests showed that they chose leaves in a similar state of decom- position. A survey of gut contents merely confirmed that organic detritus
formed the bulk of material ingested by both species in the field. A
subtle difference in diet, may, however, provide an alternative explanation
of the numerical dominance of K. vulgaris in the scalded litter plots in
Section 7.2 c)ii), if this modified litter was a more acceptable food source
to this species than to K. angulosus.
c) Microhabitat differences
In discussing the diversity and abundance of the soil fauna,
Ghilarov (1977) suggested that the structural complexity of the soil environ-
ment was the most powerful factor causing this phenomenon. Consideration
of the diversity of the cryptozoa must take into account the effect of the
relative size of the animals with respect to the litter and the soil spaces.
Using the arbitrary macro-mesofauna distinction and imagining the soil en-
vironment as it must appear to average-sized members of each group, we
realise that both potential habitat and potential food present very different
pictures. For animals relying on pre-existing cracks and pores, the
available habitat extends much deeper for the mesofauna; small spaces must 244 present a far. greater diversity of conditions for them than do the large crevices occupied by both groups. Again, the mere size of mouthparts of the macrofauna limits the amount of feeding selectivity possible, whereas mesofaunal species are able to graze fungal hyphae or bacterial colonies individually. Macrofaunal species can cover more ground and select food in a different way, but the grain of their environment must be much finer than that of the smaller animals. The far greater diversity of the mesofauna is not surprising, especially when the short generation time of small animals is considered (Ghilarov, 1977).
Within a closely related group, it has been shown that heterogeneity of habitat is sufficient to allow different species to coexist in a small area, each having a slightly different ideal microhabitat (Freeman, 1967,
1968). Complexity and dispersal of habitat will allow coexistence of potential competitors (Rathcke, 1976).
Partitioning of microhabitat space might be expected, if space is limited and determines soil animal numbers, as indicated by the studies of
Gill (1969) and Anderson (1975a). Wallwork (1976) points out that micro- habitat diversity has microclimatic, structural and biochemical features.
Each of these changes over vertical and horizontal gradients, and individual studies generally concentrate on one of these, as indeed has this one.
Vertical microhabitat differences
In forest soils, the vertical plane is characterised by profound environmental differences over short distances. Temperature and moisture gradients occur, and the amplitude of fluctuation of these parameters decreases as depth increases (Figure 5.4 and Wallwork, 1976). The slow downward movement of decomposing litter causes a number of structural and biochemical gradients. Through the litter, fermentation and humus layers, particle size and thus size of spaces decreases, while tannin and polyphenol content decrease and litter becomes more palatable to the cryptozoa. Degree of fungal colonization increases to a maximum in the fermentation layer, 245 where the highest faunal diversity occurs (Wallwork, 1976). Structural and biochemical diversity increases with depth, although in the humus layer the proportion of stable humic substances is high. In the soil itself, cavity size decreases with increasing depth as does the average body size of the non-burrowing fauna (Haarlov, 1955).
Differences in vertical distribution have been found to separate apparently coexisting species in many cryptozoan groups. Collembola
CRaarlov, 1955; Takeda, 1978) and both oribatid (Anderson, 1971; Lions,
1978) and mesostigmatid mites (Usher, 1971) display interspecific differ- ences. Studies of macroarthropods are less numerous, possibly because the body size of this group limits many species to the litter and litter/soil interface. Several studies of coexisting isopod species, however, have revealed vertical separation (Kato, 1976; Tsukamotu, 1977). Davis and
Sutton (1977), investigating the distribution of three isopod species and a diplopod species coexisting in dune grassland, found significant vertical separation between the diplopod and the isopods, and horizontal separation of one isopod species from the other two. They postulated that movement at certain times would reduce feeding competition between these two species.
Horizontal microhabitat differences
In woodland and forest, the heterogeneous distribution of litter thickness, fallen wood and topographical features form nicroclimatic, structural and biochemical differences similar to the vertical differences discussed above.
Brereton (1957) lists isopod species with their preferred micro- habitats in woodland. Blower (1955), O'Neill (1967) and Striganova (1975) have found spatial separation between coexisting diplopod species.
The horizonal complexity of litter habitats has been postulated
(Freeman, 1968) to provide a mosaic in which different species may dominate in adjacent areas because of limited movement between them and slightly different conditions within them. This hypothesis may be tenable with 246 respect to the less mobile groups of soil animals.
The vertical separation of K. vulgaris and K. angulosus is seen as a means of reducing competition for habitat space at Anglers
Creek. Each species appears more suitably adapted for the microhabitat in which it is found. Microhabitat segregation may indicate that space is limited, or that it has been in the past. Fighting has been observed between individuals in the laboratory, but sufficiently rarely that it is unlikely that interference competition is important in the ecology of these animals.
The lower oxygen consumption of K. angulosus compared with K. vulgaris has been mentioned above. Oxygen concentration in the soil atmosphere tends to be lower than that on the surface (Wallwork, 1970; Richards, 1974), and this, or the carbon dioxide concentration may prove to be important in causing the exclusion of K. vulgaris from the deeper soil layers. Even if
larger specimens can penetrate the smaller cavities at depth (Haarlov, 1955)
they will more quickly exhaust the available oxygen than smaller specimens, whose respiration is more likely to be in equilibrium with the replenishment rate of oxygen from the surface.
The difference in rate of oxygen consumption may thus be a mechanism
for habitat separation, causing a partition of limited space. d) Time
As .a niche dimension, time is generally partitioned by seasonal variation in activity, or differences related to time of day (Schoener, 1974).
Relating to the soil fauna, the niches of predatory species are often found to be segregated partly or wholly along the tine dimension, for instance in carabid beetles (gallwork, 1976) and spiders (Uetz, 1977; Turner and
Polis, 1979). This is often due to differences in time of daily activity,
• an effective means of partitioning a food resource if, like a mobile prey species noving out of cover, it is quickly renewed. However, the time dimension is more likely to be partitioned by detritivores on a seasonal basis. 247
The higher mean activity level of K. vulgaris probably provides a certain amount of temporal niche separation from K. angulosus. Newly exposed sources of palatable litter, as demonstrated by the colonization experiment, are probably available to the more active species. first, thus partitioning the food source and habitat space.
K. vulgaris displays some characteristics of a fugitive species
(Efutchinson, 1951), while K. angulosus appears to be K-selected in some situations. The success of a species may be measured in several different ways. If we consider the number of localities colonized, K. vulgaris is by far the more widespread (Figures 4.16 and 4.20) being found in 477 samples in Tasmania compared with 64 containing K. angulosus. On the other hand, if success is measured by population density at a given spot,
K. angulosus occasionally develops far larger populations than K. vulgaris ever does. It seems that where the conditions are right, the ability of
K. angulosus to utilise the soil layers allows it to use more habitat space than can a more litter-dwelling species like K. vulgaris, and thus build up much higher numbers. Unusually high densities developed by K. angulosus were found at the following localities:
Beside Emu R., Burnie, N.W. Tasmania
Beside Ferguson Falls, south of Burnie
Near St. Columba Falls, Pyengana, N.W. Tasmania
Elephant Pass, near St. Mary's, E. Tasmania
Several localities between Little Swanport and Tooms
Lake, including Anglers Creek
Near Triabunna, E. Tasmania
Maatsuyker I.
Near the east end of Prion Beach, south coast of Tasmania.
K. vulgaris occurred in smaller numbers at all but the last three of
these places, where it was absent.
These two measures of success by the species provide the clue to
the means by which each has persisted in the presence of the other, by 248 specialising in a mode of survival at which it surpasses the other species.
Of the four hypotheses advanced to explain the coexistence of detritivores in the litter and soil (p.237) each probably describes a means of avoiding competition between the two amphipod species at Anglers
Creek. The most important appear to be the regulation of population numbers by limited habitat space, and the partitioning of this space by the two species. In this case, differences in microhabitat reduce competition for space, rather than for food, as suggested in the original hypothesis.
Summary
Two landhopper species, Keratroides vulgaris and K. angulosus, coexist with apparent stability at Anglers Creek and in other parts of
Tasmania. K. angulosus is found at deeper levels in the soil, and morphological, behavioural and physiological evidence indicates that this species is better adapted to a subsoil existence than is K. vulgaris.
The higher level of activity, greater ability to colonize new habitats and very wide distribution in Tasmania indicate that K. vulgaris avoids competition /with K. angulosus by continually moving into new areas.
K. vulgaris grows to a much larger adult size than K. angulosus and differ- ences in activity and vertical distribution appear to be largely related to this size difference.
Habitat space at Anglers Creek is apparently limited now, or has been in the past, and it is thus partitioned by these two species in the vertical dimension as well as the horizontal and temporal dimensions.
This investigation is of a preliminary nature only, and further experiments to confirm or disprove its indications have been outlined. 249
CHAPTER EIGHT
POPULATION DYNAMICS AND REPRODUCTION
A study of the populations of Keratroides angulosus and K. vulgaris at Anglers Creek.
Section 8.1 Introduction
An understanding of population dynamics and reproductive biology is necessary to allow an appreciation of the function of a population in its ecosystem, yet despite their wide occurrence, terrestrial amphipods have been the subject of only two such published studies. Duncan (1969) provided detailed data on populations of Orchestia hurleyi Duncan and
O. patersoni (Stephensen) living in a disturbed grassland habitat at
Dunedin, New Zealand. The other study was carried out in a cedar plant- ation near Mito, Japan, by Tamura and Koseki (1974) who followed the dynamics of a population of "Orchestia platensis japonica" (designated
"Orchestia sp." by Morino, 1978) during a six-month period. Both of these studies thus involve species of the sexually dimorphic group (Chapters 2 and 4), although the two New Zealand species display sons apomorphic features of morphology. For instance, the second gnathopod of the male of
O. hurleyi is mitten-shaped, like that of the female, although the first gnathopods are still sexually dimorphic (Duncan, 1969; see Table 3.1, present work). O. patersoni has extremely reduced pleopods, and the hand of the male second gnathopod, although much heavier than that of the female, bears a large scabrous lobe and shows some evidence of development towards the mitten-shaped morphology found in the female. These two species, however, do not display the degree of apomorphy found amongst the sexually similar group, and no species of this group has been the subject of a published population study. Parallel development of morph- ological and physiological adaptations to the terrestrial environment has already been suggested (Section 4.4). It is thus possible that new 250 patterns of reproduction may also be found amongst the more morpho- logically advanced groups.
The method of reproduction found in amphipods, like that of isopods, is extremely important in alluding this group to colonise ter- restrial habitats. Whereas many crustaceans have planktonic larval stages, the eggs of amphipods are retained beneath the thorax of the female, where each embryo develops within the egg membrane. This event- ually breaks, releasing a first instar juvenile, which is essentially a miniature version of an adult. Newly hatched young remain for a short time in the marsupium, which is formed from a number of leaf-shaped epipodites, or Oostegites. Departure from and re-entry to the marsupium have been recorded in a number of marine and supralittoral species (Croker,
1968; Sheader and Chia, 1970; Williams, 1978). After final release from the mother, growth through a series of instars separated by moults, and development of sexual characteristics, bring the young amphipods to the reproductive stage.
Amphipod progeny are thus subject to a high degree of parental protection, which greatly reduces their early mortality rate, compared to crustacean young released as plankters. In the terrestrial and supralittoral Amphipoda, brooding keeps the eggs and newly hatched young in an extremely humid environment until they develop to a stage where they can keep themselves, by their own movement, under conditions of high humidity. The brood sizes of amphipods living in non-aquatic habitats are smaller than those of other species, because the eggs are much larger
(Bulycheva, 1957; Hurley, 1959). Consequently, newly released young are much larger and this, besides bestowing other benefits, reduces the surface area:volume ratio and thus the tendency to desiccate. It is to be expected that the more advanced landhopper species will display further adaptations to promote survival in the terrestrial environment.
The population studies reported here form part of a broader study of energy flow and nutrient dynamics at the Anglers Creek study site. 251
Section 8.2 Methods
The amphipod populations at Anglers Creek were sampled twenty- three times between 22 November 1975 and 2 December 1977. From the commencement of sampling until 28 April 1976, the populations were sampled non-quantitatively for assessment of their structure and the relative abundance of the species present. Samples were taken at irregular intervals (Figure 8.1a, b) from the general area shown in Figure 7.1.
Bulk samples of litter and soil to a depth of 10 cm were removed and transferred in a large plastic box (shown beside the car in Plate 5.3) to the laboratory for sorting.
On 14 July 1976 however, a programme of quantitative population sampling was initiated, involving collection of amphipods at approximately monthly intervals. This was continued until 2 December 1977. The area
B shown in Figure 7.2 was laid out, 20 m long and 10 m wide. This area was divided into two squares of side 10 m and on each sampling occasion, six 10 cm by 10 am quadrats were located in each square by use of random numbers. This stratified random sampling technique was used to even the spread of sampled quadrats within the area. Thus twelve quadrats were
chosen on each occasion, and a sample taken from each, using a soil corer with an internal diameter of 7 am. The corer was rested on the litter
surface at the chosen spot, while the litter was cut around it with a
sharp knife. The corer was then pushed 10 cm into the soil, twisted,
and carefully removed so that the friable soil did not fall out. The
contents of the corer from each sample were placed into a labelled poly-
thene bag, which was loosely tied for transport to the laboratory.
The samples were stored at 5 °C until sorting, which was generally
carried out within 48h. Extraction of amphipods from soil and litter
samples was achieved by manual sorting. Preliminary experiments with
the available Tullgren-type funnels showed that these were inefficient
(10-75%) in extracting amphipods. Duncan (1969) and Tamura and Koseki 252
(1974) found funnels to be over 95% efficient in the extraction of amphipods. The difference noted in the present study may have been due to the friability of the soil at Anglers Creek which led to the complete disruption of any structure during sampling.
The amphipods were sorted from the soil and litter and placed individually in 70% alcohol until they were dead. This allowed any eggs or young escaping from the marsupia of females to be collected and pre- served with the mother for recording. Amphipods were examined under a stereomicroscope, identified to species, and placed in size classes.
The length of each individual, from the anterior edge of the cephalon to the posterior end of the telson, was used to assign each specimen to a half-millimetre size-class.
The sex of each specimen was determined in the following way.
The presence of genital papillae (paired penes) on the ventral surface of peraeonite 7 was used to identify the males. Females were determined by the possession of obstegites on peraeopods 3-5, and the presence or absence of setae on these brood lamellae was noted. If neither penes nor oOstegites were present, the sex of the individual was designated indeter- minate. In the case of females bearing eggs or young, the number of progeny was determined, and the maximum length and developmental stage of eggs, or the length of young, was recorded. Dead eggs, which were wrinkled and opaque, were not included in brood counts. The stages of egg development used were similar to those of Williams (1978):
Stage A: Eggs small, colour not localised, bearing white spots
in some cases. Colour purple when alive, fading to
off-white in alcohol, white spots still visible.
Stage B: Colour localised into a crescent, somites
visible in later examples as ridges on outer surface
of crescent.
Stage C: Limbs visible, pigmented eye found in late stages.
The use of a corer of such small diameter meant that a proportion 253 of the amphipods were damaged. These specimens were counted only if the head was present. Keratroides angulosus and K. vulgaris, the two most common species at Anglers Creek, can be differentiated on head shape alone (see Key, Section 3.3), while K. vulgaris and Mysticotalitrus may be separated on the basis of antennal length. As a relationship between the number of flagellar segments on the second antenna and body length was demonstrated for each sex of each species, the presence of one second antenna allowed the length of an incomplete animal to be calculated, if it could be allocated to the male, female or indeterminate groups. Animals so incomplete that they could not be so allocated, but which were apparently above the minimum generally sexable size (K. vulgaris, 5.0 mm, K. angulosus,
4.5 mm) were assigned to the male or female groups according to the numbers of whole specimens in those two groups on that sampling date. The greatest percentage of a sample dealt with in this arbitrary manner was 5%, on
27 February 1976.
Manual sorting is a time-consuming exercise, and this restricted the number of samples sorted. The original intention was to use antennal segment number as an age indicator, as it has been preferred to body length by various authors, in work on other amphipod species (Cooper, 1965; Duncan,
1969; Tamura and Koseki, 1974; Morino, 1978). During analysis of early samples, it was found that use of this parameter gave a large number of population classes, and correspondingly reduced the numbers in each class.
For this reason, and because body length gives a better indication of body weight (on which a population energetics survey is to be based), length classes were employed in this population study.
As stated above, Keratroides vulgaris and K. angulosus were the most common amphipod species at the Anglers Creek site. Mysticotalitrus spp. were also found there, but in much smaller numbers, and the account below concerns only the two Keratroides species.
The frequencies of amphipods in each sex and size class were trans- 254 formed into percentages of the total number of each species collected on each sampling occasion. These percentages are represented in histograms in Figure 8.1.
The structure and floristics of the Anglers Creek site have been described in Chapter 5. It was mentioned in Chapter 7, however, that the area with a 100% canopy of Olearia argophylla and a very open ground layer was limited in extent. The population sampling area B was situated in this formation because of ease of movement within it. Areas away from the study site had a much more open canopy, and a fairly dense ground layer and understorey. A second site was therefore chosen, at which vegetation structure was more representative of the area as a whole, another 20 in x 10 m area was marked out, and a similar quantitative sampling procedure carried out on 26 October 1977. This was done in an effort to assess whether or not the amphipod population in area B was unusual in density, species com- position or structure. The diagrams labelled "Open site" in Figure 8.1a and b represent the sample taken on that occasion.
The second study site is situated approximately 0.5 km from area B
(Plate 5.1) in tall open-forest (Specht, 1970) dominated by Stringybark
Eucalyptus obliqua L'Herit., and White Gum E. viininalis Labill. The mod- erately dense understorey layer is composed largely of Stinkwood Zieria arborescens Sims, and Black Wattle Acacia mearnsii De Wild, with occasional
Musk Olearia argophylla F. Muell., Dogwood Pomaderris apetala Labill. and
Honeywood Bedfordia salicina DC. Native Currant Coprosma quadrifida
(Labill.) Robinson, Fireweed Senecio linearifolius A. Rich., Cutting Grass
Gahnia radula (R. Br.) Benth. and Austral Bracken Pteridium esculentum
(Forst.) Diels are found occasionally in this understorey, depending on the situation. Mother Shield-fern Polystichum proliferum Presl. forms very dense patches, sometimes covering the forest floor completely.
The most obvious difference between the two areas sampled is the absence from the second site of the 100% Olearia candopy, allowing the 8 56 280 174 67 228 125 146
1
\‘ I
2 22.xi.1975 12+1976 1.ii 1976 27.11.1976 13 iv M .1v .1rM 14.v11.1976 11.v1ii.1976 •
8 106 105 126 215 124 154 111 111 • U 26 • = ••••••••
4 MEN
2 8.60976 8x 1976 10 .19M 8.xii-1976 4.1.1977 9.11.1977 91111977 14.1v-1977
8 115 236 134 153 91 87
••1 mom C '3 4 moumnim 1111•N. M111 MI& Open 2 N.19n ' 8.'4.1977 evil 1977 17.,411.1977 191x.1977 ulorn 2.xii.1977 site 26.x 1977
I , 4 0 5 10 15 20% Percentage frequency
Figure 8.1a Percentage frequencies of male and female K. angulosus in 1/2mm length classes at Anglers Creek site on each sampling occasion. See Figure 8.1b for full explanation.
is • 48 • 85 1 17 60 71 30 1 1 t. 10 OEM 1=1= =MEIN IMO ME= =I= MUNN N 5 MI \\N. NN. N N N.\ — \\\\\\\ 10 EL NN
22•xi•1975 12-1-1976 110976 TH(1976 • 281(11976 110,41976 11-viit19M
15
39 49 47 34 96 98 •71
1.1.1 1111 NMI 11 N, 11 NN 111111111111NM 1111.a\ NX ON Ns\ \\.
81)(1976 111(1976 10-sM76 •8-xii•1976 1..(1977 91i- 1977 141v1977
59 84 49 68 34 66 112 46
IOU =MO= M&. ' Open 10.w1977 8.vi•1977 8.vii•1977 17-viii' 1977 19-b(19n 261(19n 21d(1977 site 2610977
I 0 5 10 15 20% Percentage frequency
Figure 8.1b Percentage frequencies of male and female K. vulgaris in 1 mm length classes at Anglers Creek site on each sampling occasion. Males are shown to the left, and females to the right of each diagram. Solid shading represents ovigerous females, and diagonal hatching, animals of indeterminate sex. The total number of animals in each sample appears at the upper left of each diagram. 255 development of a lower, more diverse understorey and the penetration of more light to the ground fern layer. The litter layer here is 3-5 cm deep, and the soil water content on 26 October 1977 was 39.3±4.0%, much lower than that at the main study site, which was 52.2+5.2%.
Section 8.3 Results
Population density
The mean density of both species varied considerably over the period of quantitative population study (Figure 8.2). The maximum and 2 2 minimum densities of K. angulosus were 8165/m and 1953/m , while for 2 2 K. vulgaris these values were 2431/m and 651/m , respectively. Periods of highest density occurred during summer in both species and corresponded
to the .recruitment of immature animals into the populations. The lowest
densities of both species occurred during the winter months. The large
variation of the mean density between sampling dates, however, may be partly explained by the variation between individual samples on the same
occasion. When the densities of each species on consecutive dates are
compared by a difference of means t-test, no differences are significant
except those between samples collected on 26 October 1977 and 2 December
1977 (for both species, at the p=0.05 level of significance).
It is interesting to note that the densities of both species at
the "open site" sampled on 26 October 1977 were very similar to those at
the .main study site.
Population structure and individual growth
The changes in population structure of both species during the
two years of sampling are shown in Figure 8.1a,b. K. vulgaris individuals
grow to a far larger size than K. angulosus individuals although they are
only slightly larger on release from the marsupium. This species there-
fore passes through many more half-millimetre size classes than K. angulosus.
As K. vulgaris was the less numerous species in samples, the number of 6000-
5000 -
2000 -
1000-
A S 0 A M ' J 1 J 'A 1976 1977
Figure 8.2 Densities of K. angulosus and K. vulgaris at Anglers Creek site on each sampling occasion during period of quantitative sampling. Vertical bars indicate standard deviations. 256 size classes used was reduced by using millimetre divisions in
Figure 8.1.
The populations of both species contained males, females and small animals of indeterminate sex during the entire period studied.
Ovigerous females, on the other hand, were virtually absent during the colder months of April through to August. The seasonal nature of reproduction which is apparently found in both species, would suggest that cohorts of young animals are produced annually. It might be expected therefore that their movement through the size classes could provide a useful means of estimating rates of growth of individuals.
(a) K. angulosus
Figure 8.1a shows that most size classes of males, females and unsexable animals are represented throughout the year. It is also apparent that there are no well-defined peaks which may be followed through the year to determine growth rates. While there are times when a pulse of animals in the smallest size classes (2.0-2.5 mm, and 2.5-3.0 mm), enters the population (on 1 February 1976, 8 December 1976 and 2 December
1977), there are small animals in the population continuously between these dates and the peak is thus hard to define. However, the period of each year when ovigerous females are present is much longer than the period when none are found (Figure 8.1a) so it is reasonable to expect that the corresponding dearth of young entering the population will show up more clearly than the broad peak resulting from the long breeding period.
In fact, on the female side of the histograms there is a trough which may be followed through the size classes, caused by the growth of a residual group of small animals from the previous breeding season.
These animals apparently grow when temperatures rise in early summer, moving out of the small size classes, but not being replaced for some time by the new season's hatchlings. This trough may be followed through 257
the size classes to give the rate of growth of the females even though
it does not correspond directly to a cohort of animals. The phenomenon
may be seen in each year's histograms, and Figure 8.3 shows the movement
of each trough. These data suggest that the largest females have re-
mained in the population for about two years.
By the end of the summer one year after its origin, the trough
being followed is amongst the mature female size classes. This indicates
- that at least some of the one-year females breed in their first summer,
and thus two generations of breeding females are present. There is very
little growth of the females apparent between May and September each year.
This is the time when the ecological mean temperature is below 7 °C (Figure
8.4). The overwintering female population includes two generations of
females, one immature and the other having passed through one breeding
period already.
The growth of the males is more difficult to follow. When the
histograms in Figure 8.1a are plotted separately for males, it is possible
to see the effect of the new recruitment into the population which shows
up as an increase of small males after February. At this time, a large
proportion of the overwintered male population is still present. As males
are apparently able to produce spermatozoa very soon after they are
sexable (see section on Breeding, below) this one-year cohort must make
up part of the breeding population during summer. The male population
is only strongly biased towards one size class in January and February,
so it is probable that the oldest males live only until early summer
of their second year, being available for reproduction then but disappear-
ing from the population before the maturation of the male progeny of
that breeding season.
These results indicate, therefore, that K. angulosus has a two
year life cycle, with the oldest females probably surviving two or three
months longer than the oldest males. This difference in longevity is
12 10 K.vulgaris ?? 8
6 I I I E 4
K.angulosus 99 7
6
5
4
3
2
ND J AMJJA 0 FMAMJJ 0 1975 1976 1977
Figure 8.3 Size classes corresponding to troughs on "female" side of histograms in Figure 8.1, showing growth of individuals at Anglers Creek site during entire period of sampling. 258 not sufficient, however, to account for the observed difference in size between males and females, and it is suggested that growth rates are faster in female than in male K. angulosus.
(b) K. vulgaris
The density of this species at the study site was always much lower than that of K. angulosus, and consequently the numbers on which the histograms in Figure 8.1b are based are too low to support firm conclusions. However, a minimum amongst the female frequencies similar to that followed in K. angulosus may be traced from 4 January 1977 through to the end of sampling (Figure 8.3). This indicates that females re- cruited in summer 1976-77 reached maturity by the following summer, but were still much smaller than.the largest individuals. It appears, then, that the life cycle of this species also lasts for two years.
Breeding and development
The breeding activity of a population at different times of year- may be assessed by calculating the gravidity index. As used here, this is the proportion of the adult female population which is bearing eggs or young at a particular time. The smallest egg-bearing females of
K. angulosus and K. vulgaris were found in the 5.0-5.5 mm and 6-7 mm size classes, respectively. These sizes were therefore taken as the lower limits of the adult female populations. The variation in the gravidity index in each species during the two years of study is shown in Figure 8.4.
The lack of brooding females during late autumn and winter is quite apparent.
Breeding in both species commenced in late August and ceased during April in both 1976 and 1977. Figure 8.4 compares the variation in gravidity index with both the ecological mean temperature, calculated for each month
(Chapter 5) and the duration of daylight during the sampling period.
The proportion of K. vulgaris females brooding increased more rapidly than that of K. angulosus in spring 1976. However, this phenomenon -
+ + + + + + + + + + + - + Kmulg X X X X X X X X X X - - X X X X
-15
-14 ED -13 4t1 FEC - 12
-11 3 — K.angulosus _ 06- 10 - 9
02-
KNulgaris
+ + + + + + + + + - - — + + + + KAng X X X X - X X X X X X X X - X - X X X X I i I I T I I I I I I I I T I I I I I I I I I I I NO.1 FMAMJ J A SO NO J FM AMJ J A SO NO 1975 1976 1977
Figure 8.4 Breeding data from Anglers Creek site. Variation in gravidity index (see text for definition) of K. angulosus and K. vulgaris populations at Anglers Creek site during entire period of sampling. Daylength and ecological mean temperature at the soil/litter interface (from Chapter 5). Gravidity index : K. angulosus - solid line joining dots, K. vulgaris - broken line joining dots. Daylength: smooth curve. " Ecological mean temperature: stepped line. (+) and (x) show dates on which females with setose oostegites and males with extruded spermatozoa, respectively, were found in the populations (K. vu1garis, above, •K. angulosus below figure)„ (-) indicates the absence of animals in each of these conditions. 259 was not repeated in 1977, so it appears that the breeding periods of these two species are not permanently out of phase.
Records of lengths of marsupium young showed that the largest were in the size class 3.5-4.0 mm in K. angulosus and 4.5-5.0 mm in
K. vulgaris. Incubation time is indicated approximately by the time between the spring increase in gravidity index and an increase in the density of young in the smallest size classes (Figure. 8.5). K. angulosus young below 3.5 mm in length and K. vulgaris young below 4.0 mm were chosen to indicate this increase. As the increases in brooding females and young each occur over a period, rather than at a single date, and because only monthly samples were taken, a maximum and a minimum incubation time only may be determined this way. Table 8.1 shows these ranges for each species in each year.
TABLE 8.1 Incubation time of K. angulosus and K. vulgaris during
the period of study (days).
1976 1977
K. angulosus 28-88 37-107
K. vulgaris 118-182 37-107
On hatching, the young are retained in the marsupium for a short time. While in the marsupium, young of K. vulgaris bear long plumose setae on the pleopods (which are reduced to vestigial stumps in adults). These are not found in K. angulosus young (Friend, 1979).
Soon after release, however, K. vulgaris young apparently lose these setae, as they are rarely found in free animals, even amongst the smallest individuals. The first appearance of animals bearing long pleopod setae after the non-breeding period is shown in Figure 8.5.
The sequence of development is summarised in Table 8.2 which is explained below. K.angulosus (<3.5mm )
1000
500 (.7
In 1000 CD K. vulgaris ( (4.0 mm
500
A 0 A
Figure 8.5 Densities of young in K. angulosus and K. vulgaris populations at Anglers Creek site during period of study. "Young", here, are animals below 3.5 mm (K. angulosus) or 4.0 mm (K. vulgaris) in length. 260
TABLE 8.2. Development of sexual features and growth in
K. angulosus and K. vulgaris at Anglers Creek.
Size ranges in mm.
K. angulosus K. vulgaris
size class size class
Smallest animal in marsupium 2.5-3.0 3.5-4.0
Range of marsupium young 2.5-3.5 3.5-4.5
(9 broods) (4 broods)
Smallest free animal 2.0-2.5 2.0-2.5
Largest animal with setose pleopods - 4.5-5.0
Largest animal of indeterminate sex 6.5-7.0 8.0-8.5
Smallest d' with penes 2.5-3.0 3.0-3.5
Smallest d' with spermatozoa 3.0-3.5 4.5-5.0
Smallest 9 with obstegite primordia 3.5-4.0 3.0-3.5
Smallest 9 with setose obstegites 5.0-5.5 6.0-6.5
Smallest 9 with eggs 5.0-5.5 6.0-6.5
Largest y with non-setose obstegites 8.0-8.5 12.0-13.0
Largest cr 6.5-7.0 12.0-13.0
Largest 9 9.0-9.5 15.0-16.0
The appearance of oostegites is the first indication that an
individual is female. This apparently occurs over a wide size range,
as may be seen by comparing the sizes of the smallest females with the
largest animals of indeterminate sex. The appearance of setae on the
apices of the obstegites apparently indicates that the female is in
breeding condition, as during the months when no ovigerous females are
found, there are usually no females with setose obstegites (Figure 8.4).
Also, the smallest females with obstegites bearing setae are no smaller
than the smallest ovigerous females (Table 8.2), suggesting that this
condition only just precedes the production of eggs. As the largest 261 females have been through two breeding periods and as no females have setose oostegites in at least some winter months, it follows that females must lose these setae between the two breeding periods.
In both species, the smallest sexable males are smaller than the smallest sexable females (Table 8.2). Like the females, males are probably not sexually mature until some time after the first appearance of sexual features. Sometimes preserved males are found with a silvery string attached to the penes. Examination of this material has proved that it is a mass of spermatozoa, although this may be ejected during preservation. The presence of this material indicates sexual maturity and indeed, the smallest males found with a spermatozoa mass are larger than the smallest animals with penes (Table 8.2). It appears that the production of spermatozoa is interrupted during the non-breeding season.
The only samples in which males with spermatozoa were not found were taken during the non-breeding months (Figure 8.4). Rather than being an artifact of preservation, it is perhaps more likely that this spermatic material is extruded while the male is still alive, in response to the presence of females in breeding condition. This would explain its seasonal occurrence.
Table 8.3 summarises the data relating brood size to the length class of the mother. This analysis is somewhat hampered by small sample size in the smaller and larger size classes. While the maximum brood size shows some increase with female length in both species, a clear increase in average brood size is apparent only in. K. angulosus. Other workers (Duncan, 1969; Sheader and Chia, 1970; Williams, 1978) have reported that brood size decreases with advancing egg development, as the total egg volume exceeds the capacity of the marsupium. Table 8.4 shows the mean size of broods at each of the developmental stages described earlier.
Data presented apply to the period 14 July 1976 to 2 December 1977, and the results of difference of means t-tests comparing the average brood sizes 262
TABLE 8.3 Average and maximum brood sizes found in K. angulosus
and K. vulgaris females at Anglers Creek. Bracketed
figures refer to K. vulgaris females brooding eggs at
stages A and B only.
K. angulosus K. vulgaris
9 size Average Maximum n 9 size Average Maximum n class Urm0 brood brood class (mm) brood brood.
size size size size
5.0-5.5 2.0 2 4 6-7 4.0(4.0) 4 1(1)
5.5-6.0 1.7 3 30 7-8 2.0(2.0) 2 3(2)
6.0-6.5 2.0 4 34 8-9 2.6(3.0) 7 15(8)
6.5-7.0 2.4 4 39 9-10 3.8(4.3) 8 18(9)
7.0-7.5 3.2 6 31 10-11 2.9(3.6) 7 13(5)
7.5-8.0 2.9 5 15 11-12 2.7(4.3) 10 15(6)
8.0-8.5 4.4 7 12 12-13 3.7(3.0) 4 3(2)
8.5-9.0 - 0 13-14 4.8(5.7) 7 4(3)
9.0-9.5 2.5 3 2 14-15 1.0(-) 1 1(0)
15-16 6.0(6.0) 6 1(1) are shown below. The only significant decrease in brood size is shown between the last egg stage and retained young in K. angulosus. This may be due to the release of young over a period rather than all at once.
An apparent decrease of brood size between stages B and C in K. vulgaris is not significant at these small sample sizes. If the size of
K. vulgaris broods at stages A and B is compared with female size (Table 8.3) it is seen that while broods are, on average, larger, they do not increase markedly in size with the length of the female. If it is assumed that the largest brood size found in a given size class (in a sufficiently large sample) indicates the capacity of the marsupium belonging to a female of that size class, then it appears that the average brood size of K. angulosus
263
TABLE 8.4 Mean brood size (±S.D.) at each stage of development
in K. angulosus and K. vulgaris at Anglers Creek, compared
by t-test. "Y" refers to hatchlings retained in the
marsupium.
K. angulosus K. vulgaris
Stage Mean brood size n Mean brood size
A 2.61+1.29 76 3.62+2.09 21
B 2.37±1.40 30 3.83±2.48 18
C 2.76±1.40 45 2.94+1.44 16
Y 1.38±0.74 8 2.25±1.50 4
A-B : d.f. = 104, t = 0.8411-s. A-B : d.f. = 37, t = 0.2911.s. B-C : d.f. = 73, t = 1.1811.s. B-C : d.f. = 32, t = 1.2711.s. ** C-Y : d.f. = 51, t = 2.71 C-Y : d.f. = 18, t = 0.8511s. " n.s. t =133n.s. A-C : d.f. = 119, t = 0.60 A-C : d.f. = 36, n.s. B-Y : d.f. = 36, t = 1.92 B-Y : d.f. = 20, t = 1.2111.s.
approaches this capacity more closely than does that of K. vulgaris.
Eggs of both species show a substantial increase in size with
advancing development stage (Table 8.5) . . If any real loss is occurring
from late stage K. vulgaris broods, this is most likely to be the explanat-
ion. In the case of K. angulosus, it must be concluded either that the
marsupia of the females are not filled to capacity initially, or that
their capacity increases during egg development.
TABLE 8.5. Mean length (greatest diameter) of eggs of K. angulosus and
K. vulgaris,± S.D. , at each stage of development.
K. angulosus K. vulgaris
Stage Length (mm) n Length (mm) n
A 0.78±0.11 185 0.92±0.16 72
B 0.91±0.12 64 1.01±0.04 69
C 1.09±0.13 120 1.21±0.17 47 264
TABLE 8.6 Frequencies of broods at different developmental stages
in K. angulosus and K. vulgaris samples from Anglers
Creek, at each sampling date.
K. angulosus K. vulgaris
Date ABCY ABCY
14.vii.1976 2
11.viii.1976 1
8.ix.1976 2 1 2 2
8.x.1976 4 1 3 2 5
10.xi.1976 12 3 1 2 2
8.xii.1976 6 4 9 2 2 1
4.i.1977 12 6 5 1 5 1
9.ii.1977 8 3 4 3 1 3 1
9.iii.1977 4 2 2 2 2 2
14.iv.1977 2
10.v.1977
8.vi.1977
8.vii.1977
17.vi-i.1977 1 1
19.ix.1977 4 1 1
26.x.1977 8 1 2
2.xii.1977 11 4 16 4 2 2 2 1
26.x.1977 4 4 7 1 1 2 (open site)
Table 8.6 shows the number of broods of each species found at each developmental stage during the sampling period. This provides another estimate of incubation time. While the earliest broods were probably carried from August until November or December before release, broods contributing to the stage A peak appearing in the K. angulosus population 265 on 10 November 1976, had apparently been released by 4 January 1977. This corresponds to an incubation and hatchling retention time of four to eight weeks (28-56 days) at the height of the breeding season. A similar time for
K. vulgaris can be inferred from the data in Table 8.6.
Two peaks in the number of stage A broods of K. angulosus are indicated in the 1976-77 breeding season data (Table 8.6). If the average length
(size class midpoint) of females with these broods up to and on 10 November 1976
(7.07±0.51 mm, n = 19) is compared with the average length of such females on
8 December 1976 and 4 January 1977 (6.50±0.96 mm, n = 18), a significant dif- ference is found (d.f. = 35, t = 2.26 ). The early peak is apparently in- fluenced by the larger females which have already bred the previous year,and are capable of breeding in spring. The second peak then, consists of the previous year's juveniles, which start to breed later in summer after an early period of growth and maturation under a warm temperature regime.
At the open site on 26 October 1977, the K. angulosus population seems to be at a more advanced state of breeding than at the main study site (Table 8.6). This indication is supported by the appearance of very small animals in the population on that date at the open site, whereas these are not found at the main study site until the next sampling date (Figure 8.1a).
It may be that the 100% canopy of Olearia causes a slower rise of soil temp- erature than occurs in more open areas.
Section 8.4 Discussion
The density of amphipods found at Anglers Creek on 2 December 1977 is higher than any recorded in other studies. Table 8.7 shows the maximum and minimum densities found by other workers, reported in published and unpublished work.
The increase in density which occurs in early summer each year in both populations at Anglers Creek (Figure 8.2) appears to be due to the release of young animals. Numbers decrease over autumn, apparently TABLE 8.7 Densities of terrestrial amphipods reported in population studies.
Species Maximum density Minimum density Habitat Region Source ,, 2 2 nos/.m nos./m
1 Arcitalitrus 3956 513 Subtropical rainforest N.S.W. Australia Clark (1954) sylvaticus 2 Orchestia hurleyi 2670 1230 \, Temperate grassland South I., N.Z. Duncan (1969)
O. patersoni 980 320 il II II II II II
Orchestia sp. 364 76 Cedar plantation Japan Tamura & Koseki (1974)
Mysticotalitrus 389 165 Temperate eucalypt Tasmania Friend (1975) tasmaniae forest
Keratroides angulosus 6185 1888 Temperate eucalypt Tasmania Present study forest
K. vulgaris 2431 651 11 11
1. Clark chose quadrats non-randomly in clear forest floor.
2. Densities shown are estimated from Duncan's Figure 4. 267
through mortality, especially of young animals and two-year old femles.
As stated in the introduction to this Chapter, brood sizes found
amongst terrestrial amphipods are small when compared with those of aquatic
species. Talitriator eastwoodae bears 6-14 eggs (Lawrence, 1953), while
Hurley (1959) reported that broods of Talitrus sylvaticus from New Zealand
(= Arcitalitrus dorrieni, see Section 4.1) average between 3 and 4. Duncan
(1969) found the mean brood sizes of Orchestia hurleyi and 0. patersoni
females to be similar, ranging from about 3 to about 14 early stage eggs,
depending on the size of the female (his Figure 3) and between 2 and 11
late stage eggs. The Japanese landhopper, Orchestia sp. had broods
averaging between 4.8 and 5.7 (Tamura and Koseki, 1974). Another study on a species of Orchestia from Japan (Saito and Kudara, 1972) showed that
females of this species bear between 5 and 15 eggs.
By comparison, the brood sizes of the Keratroides species at
Anglers Creek are small (Table 8.3). This seems to be mainly due to the
small size of K. angulosus and to the fact that large K. vulgaris females
usually bear much smaller broods than their apparent capacity.
There are only two published studies of terrestrial amphipods in which an attempt has been made to follow life history. Duncan (1969) worked on Orchestia hurleyi and 0. patersoni in grassland near Dunedin, New Zealand.
He found that both these species had one-year life cycles and that breeding
did not occur for most of May, June, and July. Tamura and Koseki (1974)
found that the largest individuals in an Orchestia sp. population near
Mito, Japan, were just over one year old. Two sub-populations existed, - breeding in early summer and early autumn respectively, and maturing at
'different times so that little or no interbreeding occurred between the
two groups of animals. The overall breeding season thus lasted from
May until late September, with a hiatus in mid-July.
In unpublished work, Clark (1954) provided size-class frequency histograms for a population of Arcitalitrus sylvaticus at Bulli Pass,
N.S.W., stating that ovigerous females appeared throughout the year. 268
This may not have been a pure population of A. sylvaticus, however, as
A. dorrieni is also very common in this area today. Friend (1975)
found a one-year life cycle in a Mysticotalitrus tasmaniae population near Hobart, Tasmania, in which brooding females apparently occurred in all months of the year.
Published studies of supralittoral talitrid populations (e.g.
Dorsman, 1935; Bock, 1967; Morino, 1978; Williams, 1978) have also demon-
strated life cycles which are completed in one year, while breeding occurs
only during the warmer months. In the three sexually dimorphic species which have been studied, therefore (Orchestia hurleyi, 0. patersoni and
O. sp.) life-cycles similar to those displayed by supralittoral talitrid
species have been found.
The two-year life cycle found here in both K. angulosus and
K. vulgaris indicates that somatic growth of these species is slower than
that of other terrestrial amphipods, especially in K. angulosus, which is
the smallest species yet investigated. This low rate of growth cannot
be attributed merely to low temperatures during the year, because the
climate at Anglers Creek is similar to those at the study sites of both
Duncan (1969) and Friend (1975). It appears that this life cycle is an
adaptive modification of the one-year pattern which is found in many other
talitrids.
The breeding season of both Keratroides species is clearly delimited
(Figure 8.4). Both non-breeding seasons during the period of study began
in April and finished in August. This is out of phase with the annual
mean temperature cycle, but corresponds quite closely to the variation in
daylength. Winter breeding may be induced in Talitrus saltator by
exposure to a summer photoperiod regime (Williams, 1978). The non-breeding
period found by Duncan (1969) in the Dunedin terrestrial Orchestia pop-
ulations corresponds very closely to the months when daylength is less
than 9.8h (his Figure 1). Photoperiod control of breeding has been found
in other crustaceans, for example Daphnia pulex (Stross and Hill, 1968) 269 and Orconectes nais (Armitage et a/., 1973). It is suggested here that the "resting period" (Steele, 1967) of K. angulosus and K. vulgaris is controlled by daylength, and occurs when this is less than 10-11h
(Figure 8.4).
During the resting period, females of both Keratrcddes species have no oOstegite setae. This feature has previously been recorded in amphipod species displaying a non-breeding season (Steele, 1967; Morino,
1978; Williams, 1978), and is apparently characteristic of the first moult of the non-breeding period. During the breeding season, many females, including large ones, are found without these setae. As indicated earlier, the first peak of. young of the summer are generally released by January, so it would be possible for females to produce another brood before April.
Many females of all sizes are found throughout the breeding period, however, with non-setose oostegites. This, together with the fact that no more than
60% of mature females are ovigerous at the same time (Figure 8.4), supports the proposal that females bear only one brood each summer, and subsequently lose their obstegite setae, perhaps at the next moult. At the beginning of the resting period, all the females with setose obstegites apparently moult, acquiring non-setose oostegites. Table 8.6 indicates that there are still some stage A eggs being brooded in March. Bearing in mind the falling temperatures it is apparent that these eggs would have little chance of hatching before the resting period, and are probably lost during a moult when this period commences.
Mysticotalitrus tasmaniae females acquire setose . o6stegites before bearing their first brood of eggs (at about 9.5 mm body length) and retain these for the rest of their lives (unpublished data). As stated earlier, ovigerous females occur throughout the year. During the winter months, the gravidity index remains stable or increases while the density of first instar young decreases. This is apparently because breeding continues during winter, but low temperatures retard egg development, and the young 270 moving out of the smallest size classes are not replaced by new hatchlings
(Friend, 1975). The presence of overwintering broods allows .a large release of young when temperatures rise in spring, with most of summer remaining in which to produce another brood. Thus two age-groups of
M. tasmaniae are produced annually and persist throughout the year.
Two peaks of breeding activity during a life cycle must be ad- vantageous in terms of the number of young produced, especially in ter- restrial and supralittoral amphipod species, which bear relatively small broods of large eggs. . Orchestia sp. (Tamura and Koseki, 1974), M. tasmaniae, and the two Keratroides species all practise this strategy, albeit by different means. Bimodal breeding activity has also been found in a number of studies of supralittoral talitrids (Dorsman, 1935; Bock, 1967; Nagata,
1966; Morino, 1978). The additional advantage of a winter resting period is that while young are released only in warm months, a large proportion of the population does not remain in the vulnerable small body size range for a long period during the year.
The advantages of two breeding periods'in a female lifetime are fairly obvious, but there are a number of differences between the repro- ductive strategies of K. angulosus and K. vulgaris, partly resulting from
the differences in body size between them. K. vulgaris females, being larger on average, have the capacity to brood more eggs than K. angulosus females, but this advantage is offset by several factors. Firstly,
K. vulgaris eggs are larger, and this probably gives the newly released young a better chance of survival. Consequently, however, the average brood size of K. vulgaris females, even of early stage eggs, is smaller than that of K. angulosus females of the same size (compare 8.0-8.5 mm and 8-9 mm size classes in Table 8.3). Secondly, the brooding efficiency of K. vulgaris is poorer than that of the smaller species, apparently resulting in a loss of eggs during brooding. 271
Wildish (1979) has postulated that a decrease in growth rate has accompanied movement from supralittoral to terrestrial habitats in the Talitridae. Certainly most terrestrial species are smaller than many supralittoral species, and in fact a number of plesiomorphic ter-
restrial species, for example Orchestiella spp. and some sexually di- morphic New Zealand species (Hurley, 1957) are very small. The difference
in growth rate between M. tasmaniae and these two Keratroides species, how-
ever, appears to be a result of two different reproductive strategies to
overcome the effect of a short warm season and is of little relevance
to the discussion of newly adapted terrestrial species.
In fact, the array of patterns of reproduction amongst terrestrial
amphipods is likely to be rather diverse. The sexually similar South
Australian species Austrotroides crenatus Friend (in MS, see Appendix)
appears to have a one-year life cycle, but breeding occurs only in the
cooler months. This species is found in an area with a Mediterranean
climate in which summer is dry, and the mild, moist winter apparently
provides the most suitable conditions for breeding and growth. This
variety of reproductive strategies is further evidence of the long evolut-
ionary history of the terrestrial Amphipoda. 272
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APPENDICES
Appendix I Collection details and identifications of
Tasmanian material listed in Chapter 3.
Appendix II Geographical locations and identifications of
specimens of Talitridae found in fully terrestrial
situations and reported in published work.
Appendix III Supporting papers.
(a) Friend J.A. and Richardson, A.M.M. (1977). A
preliminary study of niche partition in two
Tasmanian terrestrial amphipod species. In
U. Lohm and T. Persson (eds) "Soil Organisms
as Components, of Ecosystems". Ecological
Bulletins (Stockholm), 25:24-35.
(b) Friend, J.A. (1979). Two new terrestrial species
of Talitrus (Amphipoda:Talitridae) from Tasmania.
Papers and Proceedings of the Royal Society of
Tasmania, 113:85-98.
(c) Friend, J.A. (in MS). New terrestrial amphipods
(Amphipoda:Talitridae) from Australian forests.
302
APPENDIX I
Collection details and identifications of Tasmanian material listed
in Chapter 3.
The prefix of each sample number refers to the relevant
Tasmap 1:100 000 series sheet number. Species found in each sample
are identified by a code number, and a key to this code may be found
at the end of this appendix, with a list of collectors' names.
Abbreviations etc.
TM - Tasmanian Museum and Art Gallery
A road - logging roads on Tasmanian Pulp and Forest Holdings P/L
M road timber concessions in E. Tasmania
M28
M16
T03, T08 - logging coups near Tooms Lake, E. Tasmania.
Warra 3, 4, 5, 6 - logging coups near Geeveston, S. Tasmania. River Transect 1 etc. .,-, During the Lower Gordon AScientific Survey, tracks were
cut for access and to provide study transects. These
tracks led from the banks of several rivers in Lower
Gordon R. area, roughly at right angles to the river.
Locations on each track are identified here by the code
number of the transect and the distance from the river
. bank. The map reference of the point where each track
meets the river is given here. 303
Sample No. Locality and Habitat Description Date Collector Species
7719-1 Near Loorana, King Island. 19 -iv -1976 GE 3,10 Under teatree beside creek.
7719-2 Currie, King Island. 200 yards 19-iv-1976 GE 13 from sea, beneath regrowth eucalypts.
7719-3 Near Currie, King Island. 19 -iv -1976 GE 3,10 Beside Porky Creek. Under teatrees.
7815-1 Salmon R. Leenson Rd. 26-viii-1974 AMMR 12
7815-2 Salmon R. Chatlee Rd. 26-viii-1974 AMMR 12
7815-3 Salmon R. Rd. 26-viii-1974 AMMR 12
7815-4 Salmon R. Lerunna Rd. 26-viii-1974 AMMR 12
7815-5 Salmon R. Gravel Link Rd. 26-viii-1974 AMMR 12
7815-6 Salmon R. Gravel Link Rd. 27-viii-1974 JM, LH 12
7815-7 Salmon R. Gravel Link Rd. 19-iii-1975 JLH et a/.12
7815-8 Salmon R. Lerunna Loop 27-viii-1974 JM, LH 12
7815-9 Salmon R. Lerunna Loop 19-iii-1975 JLH 12
7815-10 Salmon R. ,Salmon R. Block 27-viii-1974 JM, LH 3,12
7815-11 Salmon R., Salmon R. Block 19-iii-1975 JLH et a/.3,12
7815-12 Salmon R. Chatlee Rd. 27-viii-1974 JM et al. 3,12
7815-13 Salmon R. Chatlee Rd. . 29-xi-1974 JLH, JM 12
7815-14 Salmon R. Leenson site 27-iii-1974 JM, LH 1,12
7815-15 Salmon R. Leenson site 29-xi-1974 JLH, AMMR 12
7815-16 Salmon R. Leenson site 19-iii-1975 JLH et a/.12
7815-17 Balfour Rd., 3.7 km along 19/21-ii-1978 AMMR 12,15 road.
7815-18 Dawson Rd., crossing road 19/21-ii-1978 AMMR 12 south of Tema.
7815-19 Stephen's Rivulet, at 4 -ii -1976 RN 1,12 Arthur R.
7816-1 Hunter Island. Leptospermum 23-v-1974 JAF 3 lanigerum swamp near homestead. In litter.
304
Sample No. Locality and Habitat Description Date Collector Species
7816-2 Hunter Island. On inland dunes 23-v-1974 JAF 3 2 mi.south of homestead. In litter.
7816-3 Hunter Island. In paddock SW of 23-v-1974 JAF 13 homestead.
7816-4 At end of road to Robbins Is. In 9-ii-1979 JAF 13 a clearing, short grass, under rock. 7816-209868.
7816-5 West end of road to Robbins Is. 9-ii-1979 JAF 13 Just behind high water mark, under teatree logs in teatree. 7816-209868.
7816-7 Robbins Is. Near runnel crossing 10-ii-1979 JAF 10,12,13 S.E. seaside track. 7816-223881.
7816-8 Robbins Is. Beside track just 10-ii-1979 JAF 10 north of Kate's Point. Under a BanAsia in low brush (5'). 7816-233887.
7816,-9 Robbins Is. Halfway between 10-ii-1979 JAF 10,12,13 Kate's Pt. and junction. In swampy area beside creek. 7816-238894.
7816-10 Robbins Is. Near junction of 10 -ii -1979 JAF 10,13 main road and Brooks Ck.Rd. Low scrub. 7816-256918.
7816-11 Robbins Is. In small gully near 10-ii-1979 JAF 10 first junction. 7816-240901.
7816-12 Robbins Is. 0.5 km W of Buckby 10-ii-1979 JAF 12,13 Pt. 7816-250892.
7816-13 Robbins Is. 0.5 km W of Buckby 10-ii-1979 JAF 3,12 Pt. Under litter at backshore. 7816-250892,
7816-14 West Wallaby Is. Under teatree 11-ii-1979 • JAF 13 on S. side of central rise. 7816-196893.
7816-15 At end of road to Robbins Is. 11-ii-1979 JAF 12 In swamp 150 m from shore. 7816-209868.
7816-16 At end of road to Robbins Is. 11-ii-1979 JAF 12,13 Seawards of 7816-15. 7816-209868.
7816-17 In forest below road west of 11-ii-1979 JAF 3,12,15 Christmas Hills. 7816-308701. 305
Sample No. Locality .and.Habitat.Description Date Collector Species
7816-18 Between Christmas Hills and 11-ii-1979 JAF 3,12 Brittons Swamp. In remnant rain- forest gully. 7816-295698.
7816-19 As for 7816-18. 11-ii-1979 JAF 3,12
7816-20 West point. S. of Marrawah. 11-ii-1979 JAF 13 7816-993656.
7816-21 North side of West Point Rd. 12-ii-1979 JAF 3,12,15 7816-034649.
7816-22 6 km. west of Togari, on north 12-ii-1979 JAF 3,10,12 side of road, in teatree swamp. 7816-148643.
7816-23 Hays Tier, in gully. 12-ii-1979 JAF 3,12 7816-162643.
7816-24 South west of Montagu. 29-i-1971 BG 12
7817-1 Hunter Island. In Eucalyptus 23-v-1974 JAF 10,13 litter at back of Shepherd's Bay.
7818-1 Grassy, King Island. 19 -iv -1976 GE 10
7818-3 Grassy, King Island. Under 19-iv-1976 GE 10 treeferns beside creek.
7913-1 Beside King River railway track 2a-iv-1975 JAF 4,12 at bend below first gorge. 7913-675273.
7913-2 Hogarth Falls, Strahan. Just 2-v-1975 JAF 4,12 Above falls in closed forest. 7913-630320.
7913-3 Near Loftus Hills Memorial. 27-iii-1975 JAF 12 Murchison Highway. 7913-630320.
7913-4 Sarah Island. At south end of 16-xii-1976 JAF 12 island. In litter under teatree. 7913-720056.
7913-6 Ocean Beach, near Strahan. On 29-x-1977 JAF 3,7,13 front dune, under lupins, among grass on sand.
7913-7 Ocean Beach. At road behind dune ,29 -x -1977 JAF 3,7,13 under L. lanigerum.
7913-8 Philips Island. Macquarie 27-x-1977 DC, MH 12 Harbour.
306
Sample No. Locality and Habitat Description Date Collector Species
7913-9 Philips Island. Macquarie Har- 27-x-1977 DC, MH 12 bour. Under rocks and sand in littoral zone.
7913-11 Beside Strahan-Zeehan railway 29-x-1977 JAF 3,11,12 line 2.5 miles north of Henty Bridge.
7913-12 Beside Strahan-Queenstown road. 30-x-1977 JAF 12
7914-1 Montezuma Falls. On abandoned Easter 1976 JAF 3,12 railway track before falls. 7914-734673.
7914-2 Montezuma Tramway. In pebbles 19-iv-1976 JAF, JLF 12 in creek on side branch to tramway. 7914-743675.
7914-3 Corinna. In wet, rotten log. 22-v-1973 TMW 12
7914-4 Trial Harbour. On west facing 15-x-1977 JAF, JLF 11,12 hillside beside track running north over cliff-top. 7914-487568.
7914-5 Badger River Valley, 3.7 miles 29-x-1977 JAF 4,11,12 north of Henty Crossing. In teatree beside old railway.
7914-8 1 mile from Savage River town- 7-viii-1974 RN 1,4,12 ship in dense forest above Whyte River. TM G1604, G2015, G2016.
7914-9 Savage River, near Obsidian 6-viii-1974 RN Creek in dense rainforest. TM G1601, G2014.
7915-1 Savage River pipeline road. 12-v-1975 AMMR, RBM, 12 At 8 mile peg. PS
7915-3 Savage River pipeline road. 11-v-1975 AMMR, RBM, 3,11,12 14 miles past 4 mile peg. PS
7915-4 Savage River pipeline road. 11-v-1975 AMMR, PS 3,11,12 Between 4 and 6 mile pegs.
7915-5 South bank of Arthur River. 10-xii-1973 JLH 12 Down Croles Rd. from Trowutta.
7915-6 Savage River pipeline road. 28-iii-1974 AMMR et a/.12 In disturbed rainforest at north rim of Savage River Valley.
7915-7 Holder's Road Spur. In swamp. 28-viii-1974 AMMR, BK 11
307
Sample No. Locality and Habitat Description Date Collector Species
7915-9 Julius River. Under stones and 2-vi-1976 RM . 11,12,15 rotting wood along east bank of river.
7915-10 Julius River. Under rotting 3-vi-1976 RM 11,12 myrtle bark on ground near creek SE of Depot Rd. Julius R. crossing.
7915-11 Julius River. Under rotting 3-vi-1976 RM 12 wood, in litter near junction of Sumac and Depot Roads.
7915-12 Magnet Valley. Ridgetop north 8-i-1977 RM 11,12 of creek.
7915-13 Holder River. Under log. Area 24-i-1976 RN 12 burnt April 1976.
7915-14 Arthur River. Beech forest near Feb. 1973 AMMR 1,4,12 new bridge.
7915-15 Roger River West. In Dogwood 6-ii-1973 AMMR 12,15 litter.
7915-16 Roger River West. Feb. 1973 AMR 15
7915-17 Roger River West. Feb. 1973 AMMR 12,15
7915-18 Dip River Falls. 9-ii-1973 AMMR 12
7915-19 Magnet Mine. Across stream from 28-viii-1975 JAF 12 old wall, below mine.
7915-20 Magnet Mine. Beside road to 28-viii-1975 JAF 4,12 mine on bend.
7915-21 Magnet Mine. Beside road to 28-viii-1975 JAF 11,12 mine.
7915-22 Dip River forest. 23-ii-1974 BK 12
7915-23 West Trowutta 29-ii-1974 JM et al. 12
7916-1 14 km from Smithton along Bass 2-vi-1977 RM 12 Highway.
7916-2 10 km west of Smithton, beside 11-ii-1979 JAF 3,12 Montagu Rd. In dense teatree swamp in cow paddock on south side of road. 7916-322814.
8011-3 Breaksea Island. Port Davey 4-iv-1975 DC, JF 3,7,13 (north island). Under pigface near shore. 8011-164027. 308
Sample No. Locality and Habitat Description Date Collector Species
8011-5 Earle's Point. Port Davey. Rain- 6-iv-1975 DC, JF 13 forest near beach. 8011-126087.
8012-1 Wilmot Bay. Lake Pedder. Near 8-iii-1975 JF 1,4 shore. 8012-167583.
8012-2 Banks of Albert River. 6-iii-1975 JF 1 8012-125570.
8012-3 Albert River Valley. In leaf 6-iii-1975 JF 1,4,12 litter.
8012-4 Beside Ropeway Creek below Olga 26-i-1976 JAF 4,12 Camp. In Dicksonia litter.
8012-5 Beside Ropeway Creek below Olga 26-i-1976 JAB 9,12 Camp.
8012-6 East of Olga Camp on transect I. 27-i-1976 JAF 2,4,12 100 m down from top in regrowth litter. 8012-020734.
8012-7 On transect I east of Olga Camp. 27-i-1976 JAF 4,9,12 200 m above Gordon River. In rainforest litter. 8012-020734.
8012-8 On transect I east. of Olga Camp. 27-i-1976 JAF 2,4,9,12 Under logs and nearby litter. 8012-020734.
8012-9 Beside Gordon River at 8012- 28-i-1976 JAF 12 943841. Under Huon Pine but not in its litter. About 2 m above water level.
8012-10 Pit 16 on transect 11A. In lit- 28-i-1976 JAF 2,4,12 ter. 25 m S of Gordon R., at 8012-943841.
8012-11 Transect 11A. 60 m S of Gordon 28-i-1976 JAF 4,12 R. at 8012-943841.
8012-12 By Pit 17 on transect 11A. 155 m 28-i-1976 JAF 2,4,12 S of Gordon R. at 8012-943841.
8012-13 Near Pit 17 on transect 11A. 28-i-1976 JAF 4,12 155 m S of Gordon R. at 8012-943841.
8012-14 Near Pit 18. Transect 11A. 380 m 28-i-1976 JAF 2,4,12 S of Gordon R. at 8012-943841. In leaf litter beside track.
8012-15 Pit 19. Transect 11A. 490 m S of 28-i-1976 JAB 2,4,12 Gordon R. at 8012-943841. Under overhanging log. 309
Sample No. Locality and Habitat Description Date Collector Species
8012-16 Transect 11A. 580 in S of Gordon 28-i-1976 JAF 4 R. at 8012-943841. In litter by track.
8012-17 Pit 20 on transect 11A. 670 in 28-i-1976 JAF 4 S of Gordon R. at 8012-943841.
8012-18 Transect 11A. 720 in S of Gordon 28-i-1976 JAF 1,2,4 R. at 8012-943841.
8012-19 Upper transect 11A. 750 in S of 28-i-1976 JAF Gordon R. at 8012-943841.
8012-20 On button grass plain east of 21-ii-1976 JAF 1,2,4 Olga R. Transect 7R. 330 in from Olga R. at 8012-042550.
8012-21 In band of forest east of Olga 21-ii-1976 JAF 1,4,12 River, running parallel to it. Transect 7R. 550 in from Olga R. at 8012-042550
8012-22 On button grass plain 10 metres 21-ii-1976 JAF 4 from patch of scrub. Near heli- pad. Transect 7R. 1250 in from Olga R. at 8012-042550.
8012-23 Upper Olga Valley. Transect 7R. 21-ii-1976 JAF 1,4,6, 1300 in from Olga R. at 8012- 11,12 042550.
8012-24 Transect 7R. 1445 m from Olga 21-ii-1976 JAF 4,11,12 R. at 8012-042550. Near Pit 42.
8012-25 Transect 7R. 1535 in from Olga 18-ii-1976 JAF 11 R. at 8012-042550. Beside Pit 43 in rotting fallen teatree.
8012-26 Transect 7R. 1445 in from Olga 18-ii-1976 JAF 1,4 R. at 8012-042550. Near Pit 43 east of Olga R. in Olga-Hardwood Valley.
8012-27 Transect 7R. 1470 m from Olga 18-ii-1976 JAF 4,12 R. at 8012-042550. In rain- forest just east of Pit 43.
8012-28 Transect 7R. 1470 m from Olga 18-ii-1976 JAF 12 R. at 8012-042550. Just east of Pit 43. in Gahnia clump next to 27.
8012-29 Transect 7R. 1705 in from Olga 18-ii-1976 JAF 11,12 R. at 8012-042550. Near Pit 44, in moss and wood of rotten log. 310
Sample No. Locality and Habitat Description Date Collector Species
8012-30 Transect 7R. 1705 m from Olga 18-ii-1976 JAF 4,11,12 R. at 8012-042550. Near Pit 44, in litter.
8012-31 Transect 7R. 1920 m from Olga 18-ii-1976 JAF 1,4,11,12 R. at 8012-042550. Neat Pit 45. On west side of slope of small hill on 7R, in litter.
8012-32 Transect 7R. 1925 m from Olga 18-ii-1976 JAF 1,4,9,12 R. at 8012-042550. On top of hill at Pit 45.
8012-33 Transect 7R. 1930 m from Olga 18-ii-1976 JAF 1,4,9,12 R. at 8012-042550. On east side of hill at Pit 45.
8012-34 Transect 7R. 2093 m from Olga 18-ii-1976 JAF 4,11,12 R. at 8012-042550. Just past Pit 46.
8012-35 Ropeway transect. On side of 20-ii-1976 JAF 12 hill west of Ropeway Creek, near Olga Camp. 8012-016730.
8012-36 Ropeway transect. At top of 20-ii-1976 JAF 12 hill west of Ropeway Creek. Under a board on gravel path. 8012-96730.
8012-37 Ropeway transect. Second gully 20-ii-1976 JAF 12 on ropeway. Beside creek, in rotten log. 8012-014730.
8012-38 Ropeway transect. Second gully 20-ii-1976 JAF 12 on ropeway from Olga Camp. Amongst litter and under logs. 8012-014730.
8012-39 Ropeway transect. In pure stand 20-ii-1976 JAF 9,12 of dogwood, near end of ropeway. 8012-012731.
8012-40 Transect 8L. 225 m from Gordon iii-1976 JAF 12 R. at 8012-975796. Beside creek on 8L.
8012-41 Transect 8L. 370?m from Gordon iii-1976 JAF 4,12 R. at 8012-975796.
8012-42 Transect 8L. 450 m from Gordon iii-1976 JAF 1,4,12 R. at 8012-975796.
8012-43 Transect 8L. 450 m from Gordon iii-1976 JAF 2,4,12 R. at 8012-975796. As for 8012-43 but on lower side of tree in hollow.
311
Sample No. Locality and Habitat Description. Date Collector Species
8012-44 Transect 8L. 600 m from Gordon iii-1976 JAF 2,4,12 R. at 8012-975796. In small gully crossing track.
8012-45 Transect 8L. 770 in from Gordon iii-1976 JAF 2,12 R. at 8012-975796. Just above Pit 31.
8012-46 Transect 8L. 1075 in from Gordon iii-1976 JAF 2,4 R. at 8012-975796. Hill on side of Sprent Valley, about 20 metres above valley floor.
80/2-47 Transect 8L. 1165 m from Gordon iii-l976 JAF 1,4,12 R. at 8012-975796. Teatree flood plain beside Sprent River.
8012-48 Truchanas Reserve. On hill NW of 19-iii-1976 JAF 1,4,12 bend in Denison River.
8012-49 Truchanas Reserve. On little 19-iii-1976 JAF 12 floodbank 3 m Above Gordon R. behind shingle bed beside bend.
8012-50 Truchanas Reserve. Just in from 19-iii-1976 JAF 4,11,12 shingle bed (landing spot) on west bank of Gordon River.
8012-51 Truchanas Reserve. Near bend in 19-iii-1976 JAF 4,11,12 Gordon River.
8012-52 Beside Sprent River. 200 m east iii-1976 JAF 2,4,11,12 of the end of transect 8L.
8012-53 Beside Sprent River. 200 m down- iii-1976 JAF 2,4,12 stream from the end of transect 8L.
8012-54 Transect 8L. Just above big rock iii-1976 JAF 1,4 at the Gordon R. end of transect. 8012-975796.
8012-55 Transect 8R. 250 m from Gordon 2-ii-1977 AMMR 4,12 R. at 8012-975796.
8012-56 Transect 8R. 50 in from Gordon 2-ii-1976 AMMR 12 R. at 8012-975796.
8012-57 Transect 8R. Gordon River. At 2-ii-1977 AMMR 12 top of steep bank 15 feet above present river level.
8012-58 Gordon River Above Abel Gorge. 3-ii-1977 AMMR 12 On left bank 300 feet Above river, N-facing slope.
312
Sample No. Locality and Habitat Description Date Collector Species
8012-59 As for 8012-58. S-facing river 3-ii-1977 AMMR 4,12 bank. Vegetation more closed and more moss.
8012-60 Summit of Moores Lookout, in a 16 -xii -1976 JAY, DS, 1,9,12 patch of scrub. 8012-922610. LB
8012-61 Near creek flowing into Wanderer 16-xii-1976 JAY 4 River north of Mbores Valley Camp. 8012-805662.
8012-62 In rainforest on southern slopes 16-xii-1976 JAY 1,12 of Conder . River Valley beside track, where big fallen tree has blocked route. 0.75 km from river. 8012-803678.
8012-63 At Conder River crossing, on N 16-xii-1976 DS 12 side. 8012-803684.
8012-64 Albert River Valley. 8012-133543. 6-iii-1975 DC et a/. 4
8012-65 Olegas Truchanas Huon Pine Res- 19-iii-1976 JLH 4,6,11,12 erve. 8012-150765.
8012-66 Gully beside Strathgordon Dam Rd. 25 -iv -1978 JLH, LH, 2,4,12 In moss. 8012-164668. JAS
8012-67 Gully beside Strathgordon Dam Rd. 25 -iv -1978 JLH, LH, 2,4,12 In litter. 8012-164668. JAS
8013-1 Bird River. SW side 200 m from 18-iii-1974 JAY 12 river on Kelly Basin Rd. Under logs and moss. 8013-843110.
8013-2 Bird River. NE side, 1 km from 18-iii-1974 jAF 12 river on Kelly Basin Rd. In litter. 8013-837110.
8013-3 Bird River. Beside bridge on S. 18-iii-1974 JLH 12 side on Kelly Basin Rd. 8013-843110.
8013-4 On Kelly Basin Rd. 3.4 km beyond 18-iii-1974 JAY et al. 12 Franklin River Rd. turnoff. On sunny hillside beneath large rotten log. 8013-850128.
8013-5 As for 8013-4. Beneath small 18-iii-1974 JAY et al. 12 rotten log. 8013-850128.
8013-6 As for 8013-4. Under logs. 18-iii-1974 JAF et al. 12 8013-850128.
8013-7 On Kelly Basin Rd., 5 km from 18-iii-1974 JAY et al. 12 Bird R. bridge, 2.3 km beyond Franklin R. Rd. turnoff. South- ern aspect of hill, in moss. 8013-853133.
313
Sample No. Locality and Habitat Description Date Collector Species
8013-8 As for 8013-7. In litter. 18-iii-1974 JAF et al. 2,4,12
8013-9 As for 8013-7. Under rotten logs. 18-iii-1974 JAF et al. 2,12
8013-10 Kelly Basin Rd.,0.5km S of Clay- 16-iii-1974 JAF, PN 4 ton Creek/Crotty Rd. bridge. On west bank of road. 8013-860211.
8013-11 West side of drainage bridge, 19-iii-1974 JAF et al. 4,12 6.2 miles from Lyell Highway on Crotty-Kelly Basin Rd. Ex moss. 8013-885329.
8013-12 As for 8013-11. Ex leaf litter. 19-iii-1974 JAF et al. 4,12
8013-13 As for 8013-11. In litter and 19-iii-1974 JAF et al. 12 rotten logs.
8013-14 East slope, 2 km from junction 17-iii-1974 JAF etaal. 1,4,12 of Franklin River Rd. on Kelly Basin Rd. towards Bird River. Ex litter. 8013-852133.
8013-15 As for 8013-14. Ex rotten log. 17-iii-1974 JAF et a/. 4,11,12
8013-16 Kelly Basin Rd., 2.8 km from 17-iii-1974 JAF et al. 4,9,12 Franklin River Rd. turnoff. On E side of valley, halfway up hill.Ex litter. 8013-857167.
8013-17 As for 8013-16. In rotten logs. 17-iii-1974 JAF et al. 12
8013-18 0.5 km S of Clayton Creek, on 16-iii-1974 LC, SE,GB 4,9,12 Kelly Basin Rd., east side of road. Ex litter. 8013-860211.
8013-19 On Kelly Basin Rd., 1.5 km N of 16-iii-1974 JAF et al. 1,4,12 junction with Franklin River Rd. Steep hillside on west side of Purgatory Creek Valley. Ex moss. 8013-856161.
8013-20 As 8013-19. Ex litter. 16-iii-1974 JAF et a/. 1,4
8013-21 As 8013-19. Ex rotten log. 16-iii-1974 JAF et al. 4
8013-22 Kelly Basin Rd. just N of Water- 17-iii-1974 JAF et al. 1,4,12 fall Creek, on mule track, at edge of button grass. Ex litter. 8013-852227.
8013-23 Kelly Basin Rd., 0.1 km from 18-iii-1974 JAF et a/. 4,12 junction of Franklin Rd. Steep slope and gully, under rotten logs. 8013-858146. 314
Sample No. Locality and Habitat Description Date Collector Species
8013-24 Kelly Basin Rd., Waterfall Creek, 17-iii-1974 JAF et al. 1,4,12 on track on S side of valley. Ex litter. 8013-849225.
8013-25 Halfway up Snake Peak, beside 15-iii-1974 JAF et a/. 1,4,12 track. ENE slope. Ex litter. 8012-847234.
8013-30 Darwin, Kelly Basin Rd., from 19-iii-1974 JAF et a/. 12 seepage on S slope of hill. Ex moss. 8013-856236.
8013-31 Darwin, Kelly Basin Rd., east 17-iii-1974 JLH 12 slope of hill. Ex litter. 8013-856236.
8013-32 Darwin, Kelly Basin Rd., top of 19-iii-1974 GB et a/. 12 hill. 8013-856236.
8013-33 As for 8013-30. Ex moss. 19-iii-1974 JAF et a/. 4,12
8013-34 As for 8013-32. Under logs. 19-iii-1974 JAF et a/. 4,12
8013-35 Beside Lyell Highway, between 20-iv-1976 JAF, JLF 12 Collingwood River and Franklin 'River crossing. In litter. 8013-121295.
8013-37 Frenchmans Cap area, NW and just 18-iii-1978 JAF 12 below Tahune hut. 8013-039200.
8013-38 S. side of Sharland's Peak, in 18-iii-1978 JAF 4,11,12 King Billy pine forest. 8013-054191.
8013-39 In forest on N slope below Barron 18-iii-1978 JAF 1,4,11,12 Pass. Steep hillside, near creek runnel. 8013-059188.
8013-40 Southern end of Lake Vera, in 19-iii-1978 JAF 4,12 rainforest. 8013-072189.
8013-41 Near top of Philps Lead. 19-iii-1978 JAF 4,12 8013-092184.
8013-42 West side of King River. In 13-xii-1978 LH 12 forest soil/litter. 8013-894433.
8013-43 On east bank of King River oppos- 13-xii-1978 LH 12 ite mouth of Comstock Creek. 8013-889432.
8014-1 Moxon Saddle, N. Tyndalls, beside 29-iii-1975 JAF 12 road running N from main road. 8014-813665. 315
Sample No. Locality and Habitat Description Date Collector Species
8014-2 As for 8014-1. Collected in 29-iii-1975 JAF 2,4 litter bag, Tullgren extraction.
8014-3 Moxon Saddle, N. Tyndalls, near 29-iii-1975 JAF 12 river bed, boggy area under rain- forest. In sides of tree stumps, tussocks and moss.
8014-4 North Tyndalls, on road N from 29-iii-1975 JAF 12 Henty Camp. Clearing by a creek under leatherwood. 8014-798650.
8014-5 As for 8014-4. Collected in lit- 29-iii-1975 JAF 4,12 ter bag, Tullgren extraction.
8014-6 On road north of Henty River 29 -iii -1975 JAF 12 (ryndalls) campsite. 8014-800645.
8014-7 As for 8014-6. Collected in lit- 29-iii-1975 JAF 1,12 ter bag, Tullgren extraction.
8014-8 On road north from Henty River 29-iii-1975 JAF 2,4,12 (Tyndalls) Camp, just below a side track down to river. - 8014-800640.
8014-9 Just below Henty River Camp. 29-iii-1975 JAF 1,4,12 8014-798625.
8014-10 Below Mt. Tyndall, near creek 29-iii-1975 AMMR 12 crossing Basin Lake Road. 8014-806563.
8014-11 On Luke's Knob, between Lake 28-iii-1975 JAF 12 Westwood and Lake Julia. N. Tyndalls. Between rocky out - cropS. 8014-823619.
8014-12 100 metres N of Lake Westwood. 28-iii-1975 JAF 1,12 In scrubby patch and around erratics on button grass plain. 8014-828627.
8014-13 Near Red Hills. In litter around 28-iii-1975 JAF 1 a slight rise on glacial plain. 8014-830660.
8014-14 Moxon Saddle. Beside road just 28-v-1975 JAF 4,11 out of main forest. In moss and litter.
8014-15 North Tyndalls. In small patch 28-iii-1975 JAF 12 of litter above Lake Julia. Button grass plain hillside. 8014-819619.
316
Sample No. Locality and Habitat Description Date Collector Species
8014-16 Just up from creek on the road 29-iii-1975 JAF 4,12 N of Henty Camp. 8014-800600.
8014-17 North Tyndalls, on a flat. 29-iii-1975 JAF 4,12 8014-805668.
8014-18 Beside Dove Lake, in King Billy 25-iv-1978 JAF 12 forest below Ballroom Plateau. 8014-129865.
8015-1 Beside Murchison Highway, just 27-vi-1975 JAF 12 before descending into Hellyer Gorge from north. 8015-314830.
8015-2 Vacant block next to 4 Hodgman 15-viii-1973 DC 11,12 St., Burnie.
8015-3 Willy's farm at Calder; on ridge 28-vii-1975 JAF 12 S of paddocks, near swampy patch of teatree.
8015-4 Just above Emu River bridge (N. 28-vii-1975 JAF 12,15 side)., Burnie. 8015-095517.
8015-5 Between Fern Glade Rd. and Emu 28-vii-1975 JAF 4,12,15 River, Burnie. 8015-094521.
8015-6 David Coleman's garden, Myrtle 25-viii-1975 JAF 12,15 Crescent, Burnie. 8015-093521.
8015-7 David Coleman's grandmother's 25-viii-1975 JAF 12,15 garden, 4 Hodgman St., Burnie. 8015-077547.
8015-8 Beside road near Highdene (Mount 25-viii-1975 JAF, RN 15 Rd). 8015-996380.
8015-9 Just down 18-mile Rd., off Mount 25-viii-1975 JAF 12 Rd., behind Burnie. 8015-979342.
8015-10 Beside Ferguson Falls, Talunah 25-viii-1975 JAF 12,15 Ed. 8015-923313.
8015-11 Bush near Douglas Brook, near 25-viii-1975 JAF, BM 4,11,12,15 Oonah. 8015-882330.
8015-12 First creek along Blackwell's 26-viii-1975 JAF 12 Rd., (Inglis River). 8015-823355.
8015-13 Near crossing of second creek 26-viii-1975 JAF 11,12 on Blackwell Rd. (N. of Hellyer Gorge) in big bend. 8015-807352.
8015-14 First or second creek valley 26-viii-1975 JAF 12,15 back from junction of Arthur and Hellyer Rivers. 8015-760356.
317
Sample No. Locality.and.Habitat.Description. .Date.. Collector Species
8015-15 0.5 km up gully beside bridge 26-viii-1975 JAF 11,12 over Hellyer. River on Blackwell Rd. 8015-760356.
8015-16 Beside Murchison Highway, on 27-viii-1975 JAF 12 west side of bridge over Wandle River. 8015-813197.
8015-17 As for 8015-16 but further from 27-viii-1975 JAF 12 bridge.
8015-18 West Downs, Surrey Hills. "Pure 16/17-ii-1978 AMMR 12,15 myrtle" site on APPM reserve. In treefern litter. 8015-863229.
8015-19 As for 8015-18 but under myrtle 16/17-ii-1978 AMR 12,15 litter.
8015-20 As for 8015-18 but under logs 16/17-ii-1978 AMMR 11 in distinct burrows in clay.
8015-21 West Downs, Surrey Hills. "Euc- 16/17-ii-1978 AMMR 11,12,15 alypt" site, APPM reserve. 8015-863229.
8015-22 West Downs, Surrey Hills. "Grass-16/17-11-1978 AMMR 12,15 land" site, APPM reserve. 8015-863229.
8016-1 Near Sisters Creek, just west 9 -ii -1979 JAF 12 of turnoff to Watts Siding. N side of Bass Highway. 8016-780644.
8110-1 Maatsuyker I. In soil under 9-xi-1978 GB 4,7,13 briquette dump. 8110-419669.
8110-2 Maatsuyker I. Beside haulage 16-i-1979 JAF 4,17,13 (E side), near top under teatree.
8110-3 As for 8110-2 16-i-1979 GE 7,13
8110-4 As for 8110-2 16-i-1979 GE 7
8110-5 Maatsuyker I. Further up side 16-i-1979 GE 7,9,15 of valley (5 m) from 8110-6.
8110-6 Maatsuyker I. Just below "Olym- 16-i-1979 GE 4,9,15 pic Pool" above road. 8110-415664.
8110-7 Flat Witch I. In a shallow gully 16-i-1979 JAF 4,7 on E side. In litter between muttonbird burrows. 8110-430702.
8110-8 Flat Witch I. Under Poa at side 16-i-1979 GE 4,7 of 8110-7 gully. 8110-430701.
318
Sample No. .Locality and Habitat Description Date Collector Species
8110-9 Flat Witch I. Near top of island,16-i-1979 JAF 4,7,13 under teatree and native pepper.
8110-10 Flat Witch I. As for 8110-9. 16-i-1979 GE 4,7,13
8110-11 De Witt I. In wet sclerophyll 16-i-1979 JAF 4,7,9, beside creek 150 in from shore. 12,15 8110-482732.
8110-12 De Witt I. 300 m upstream from 16-i-1979 JAF 4,9,12,15 8110-11. 8110-481730.
8110-14 De Witt I. 200 m further upstream16-i-1979 JAF 4,9,12 from 8110-12 and 13. 8110-481728.
8110-15 De Witt I. In eucalypt litter on 16-i-1979 JAF 4,6,9,12 NW-SE ridge running across N of main gully. 8110-484727.
8110-16 De Witt I. In muttonbird rookery 16-i-1979 JAF 7,9,12,13 on N side of island behind E end of bay. 8110-481733.
8110-21 Maatsuyker I. Ex litter collec- 16-i-1979 JAF 7,9,15 ted from 8110-6 site. 8110-430702.
8110-22 Flat Witch I. Ex litter collect- 16-i-1979 JAF 4,7,13 ed from 8110-9 site. 8110-428699.
8110-23 De Witt I. Bottom site. Ex lit- 16-i-1979 JAF 9,9,12,15 ter collected at 8110-11 site. 8110-482732.
8110-24 De Witt I. Ex litter collected 16-i-1979 JAF 2,4,9, about 300 in below the top of 12,15 gully leading up to steepest pArt og cliffs on SE side of is- land. 8110-485723.
8111-1 High Celery Top I. Ex rain- 16-viii-1973 JK 4 forest leaf litter. 8111-318970.
8112-1 Small patch of scrub on button- 17-v-1975 JAF 9,12 grass plain under CE of ) the Thumbs. 8112-485748.
8112-2 On Little Florentine River, be- 13-v-1975 JAF 12 side wooden planking on Adams- field track, 1.5 miles from turnoff. 8112-527714.
8112-3 East end of Gordon Gorge in 17-v-1975 JAF 12 scrub above river. 8112-479763.
8112-4 Beside Rasselas Valley track in 17-v-1975 JAF 12 Florentine Valley. 8112-512710. 319
Sample No. Locality and Habitat Description Date Collector Species
8112-5 Huon-Wedge Divide along Scotts 3-v-1973 RBM, BK 4,12 Peak Road.
8112-6 Below Centre Star, W of Denison 5-iii-1975 CTP 12 Range.
8112-7 Little Florentine River on Gor- 28-v-1975 JAF 9,12 don River road, offside of road. 8112-527680.
8112-8 Boyd Lookout, just below road. 28-v-1975 JAF 4,12 8112-478592.
8112-9 Just before McPartlan Pass. S 28-v-1975 JAF 4,12 side of road. 8112-368543.
8112-10 Beside Gordon Rd. near Trappes 28-v-1975 JAF 1,4,12 Hill (near Strathgordon). Near gate on Holley Rd. 8112-259606.
8112-11 Between Gordon Dam and Strath- 28-v-1975 JAF 4 gordon. Beside road, just above cutting (S side of road).
8112-12 Lake Rhona, upper Gordon River 20-iii-1972 APA, HDB 4,5,11,12 area, below Reed's Peak.
8112-13 Lake Pedder. Ex vegetation 3-v-1972 4 behind sand dunes.
8112-14 On peninsula jutting into Windy 14-i-1977 JAF 12 Lake. 8112-312913.
8112-15 Scotts Peak Rd. opp. forest Con- 9-viii-1972 RR 4,8,12 servation Area 703, Lookout No.I. Ex moss.
8112-16 Banks of Huon R. on W side of 3-v-1973 JLH et a/. 4,12 bridge over Huon R., Scott's Peak Road.
8112-17 Divide between Huon and Floren- 3-v-1973 JLH 4,12 tine Rivers, Scott's Peak Rd. Ex moss.
8112-18 Florentine Valley. Under logs 26-ix-1964 JJG 12 in rainforest.
8112-19 HEC Nature trail beside Scott's 15-iv-1973 BJS, AJD 4,12 Peak Road.
8113-1 6 miles along Butlers Gorge Rd. 3-v-1975 JAF 9,12 from turnoff, over canal. 8113-443193.
8113-2 Tarraleah 7-x-1957 VVH 8,12 320
Sample No. Locality and Habitat Description Date Collector Species
8113-3 Cuvier Valley, Cradle Mountain- 29-ix-1973 AMMR, RM 12 Lake St. Clair National Park.
8113-4 On the W foothills of Algonkian 8-i-1977 JAF 4,12 Mountain. 8113-199070.
8113-5 At campsite on Algonkian Mount- 9-i-1977 JAF 4,12 am, on a ridge running towards the summit, on knoll on side of ridge, above rainforest. 8113-213062.
8113-6 In a gully near the top of 9-i-1977 JAF 4,12 Algonkian Mountain. 8113-218059.
8113-7 Near Creek south of ridge run- 9-i-1977 JAF 4,12 ning down from Algonkian summit towards Prince of Wales range. 8113-235046.
8113-8 On belt of rainforest in gully 11-i-1977 JAF 4,12 South of Diamond Peak. 8113-217981.
8113-9 Beside a creek in corridor of 12-i-1977 JAF 12 teatree and Banksia just before lead up onto east wall of Den- ison Valley at "Tilted Valley". 8113-277293.
8113-10 At campsite in "Tilted Valley". 13-i-1977 JAF 4 8113-305977.
8113-11 South side of Lyell Highway just 19-iii-1974 JLH et a/. 4,12 E of King William Saddle. 8113-278261.
8113-12 As for 8113-11. 19-iii-1974 JAF 1,4,12
8113-13 As for 8013-26. Ex foliage of 19-iii-1974 JAF 4,12 fallen King Billy pine branch. 8113-278261.
8113-14 As for 8013-26. Under log. 19-iii-1974 JLH et al. 12 8113-278261.
8113-15 East side of Franklin River on 29-x-1977 JAF 9,12 Lyell Highway. 8113-192257.
8115-1 West of the Gnomon, Dial Range. 26-viii-1974 BK, AMMR, 12 Beside second creek beyond farm, DC from Penguin. ,8115-186448.
8115-2 Gully beside small creek at W 25-iv-1978 JAF 12 end of bridge over Forth R., below Cethana Dam. 8115-276083. 321
Sample No. Locality and Habitat Description Date Collector Species
8115-3 Turners Beach, beside estuary of 12-ii-1979 JAF 12,13 Forth River (west bank).
8210-1 Near Catamaran Fiver, other side 23-viii-1979 JAF 5 of road from sea. 8210-907793.
8210-3 Rainforest gully 3 km N of Cat- 19-v-1976 AMMR 4,9,12 amaran River, above road.
8210-4 Left bank of South Cape Rivulet, 6-ix-1976 PA 5 South Coast track. 8210-826726.
8210-5 Beside creek at SW end of Blow- 26-xii-1975 JAF 4,9,12,15 hole Valley. 8210-868716.
8210-6 Campsite at South Cape Rivulet. 26-xii-1975 JAF 5,8,12,15 8210-826724.
8210-7 Beside creek 2 km past Blackhole 27-xii-1975 JAF 2,4,9 lookout, South Coast track. 8210-782722.
8210-8 In rainforest scrub just east of 28-xii-1975 JAF 4,5,6,15 Surprise Bay. 8210-724740.
8210-9 2-3 km west along track from Sur- 28-xii-1975 JAF 4,9,12 prise Bay. 8210-708757.
8210-10 1/2 mile past 8012-9 in wet sclero- 28-xii-1977 JAF 4,9,12 phyll. 8210-706760.
8210-11 Geeves campsite above eastern 29-xii-1975 JAF 15 end of Priori Beach. 8210-682786.
8210-1.3 Cockle Creek—Ex eucalypt litter. 15-vi-1972 BK, KBM, 12 PSL
8210-14 3 km N of Catamaran Beach. 19-v-1976 AMMR, BK, 7 8210-912787. DC
8210-15 2 km N of old Cockle Creek, on 19-v-1976 AMMR, BK, 4,9 new Forestry road. DC
8210-16 Forestry road over Catamaran 19-v-1976 AMMR, DC, 4,9 River, 3 km S of river. BK
8210-18 3 km N of Catamaran River, in 19-v-1976 AMMR, DC, 4,9,12 rainforest gully beside Forestry BK road.
8210-19 Beside Catamaran River at 19-v-1976 AMMR, DC, 9 Forestry road crossing. BK
8210-20 Ile du Golfe. Just over ridge on 16-i-1979 JAF, GE 7,12 E side. 8210-617759. 322
Sample No. Locality and Habitat Description Date Collector Species
8210-21 Ile du Golfe. Just above cliffs, 16-i-1979 JAF 7,12 E side, below 8210-18. 8210-618758.
8210-22 Ile du Golfe. Near 8210-19. 16-i-1979 GE 7,12 8210-618758.
8210-23 Ile du Golfe. Extracted from 16-i-1979 JAF 12 litter collected at 8110-19 site. 8210-618758.
8210-24 Near South Cape Bay, among 30 -vi -1973 JB 8,13? litter.
8211-1 On track to Kermandie from Hartz,24-viii-1973 JLH, AMMR 2,4,9,12 at head of Arve River, W of Taylor's Ridge. 8211-832142.
8211-2 On Kermandie Track to Hartz 24-viii-1973 AMMR, JLH 2,4,5,9, Mts. at head of Arve River, W 12 of Taylor's Ridge. 8211-832142.
8211-3 Hartz Mountains. Under litter at 14-viii-1975 JAF 9 end of road. 8211-873149.
8211-4 In Warra 3 coup, over Tahune 29-v-1974 JLH, JM, 9,12 Bridge. 8211-783309. AR
8211-5 Near boundary of Warra 5 and 6, 29-v-1974 JLH et al. 4,8,9,12 over Tahune Bridge. In moss. 8211-775296.
8211-6 Near boundary of Warra 5 and 6, 29-v-1974 JLH et a/. 2,4,8,9, over Tahune Bridge. In litter. 12 8211-775296.
8211-7 In Warra 4, over Tahune Bridge. 29-v-1974 JLH et a/. 4,9,12 In litter. 8211-780309.
8211-8 As for 8211-7, in moss. 29-v-1974 JLH et al. 4,8,9,12
8211-9 Kermandie track, from Hartz hut, 24-viii-1973 JLH, AMMR 2,4,5,12 at head of Arve River, west of Taylor's Ridge. 8211-829146.
8211-10 In myrtle forest beside Tahune 30 -xi -1975 JAF 9,12 Picnic area. 8211-777285.
8211-11 At hut end of old. track from 30 -xi -1975 JAF 4,9,12 lookout to hut at Hartz Mts. 8211-814150.
8211-12 Campsite at New River Lagoon end 29-xii-1975 JAF 4,9,12 of Precipitous Bluff route. 8211-659851. 323
Sample No. Locality and Habitat Description Date Collector Species
8211-13 On Precipitous Bluff - About 29-xii-1975 JAF 4,9 200' below cliffs, on ridge in low rainforest. 8211-678864.
8211-14 One hour's walk from quarry at 26-xii-1976 JAF 4,9,12 head of track to Moonlight Flats. 8211-866875.
8211-15 Scrub in flat area just before 26-xii-1976 JAF 4,9 climb onto Moonlight Flats. 8211-855883.
8211-16 Burn Creek campsite, Moonlight 27-xii-1976 JAF 4,9,12 Flats. 8211-832880.
8211-17 North side of Mt. La Perouse. 28-xii-1976 JAF 4 Under King Billy pine in a little hollow with creek. 8211-783840.
8211-18 Moonlight Flats track. At top of 29-xii-1976 JAF 2,9,12 second hill after leaving quarry. 8211-860877.
8211-19 First hill after quarry on way 29-xii-1976 JAF 4,9 to Moonlight Flats. 8211-868875.
8211-20 Hastings. Ex soil, 300 yr. 2-viii-1976 LH 9 E. obliqua.
8211-21 Hastings. 1914 block. Facy Rd. 9-viii-l976 LH 9
8211-22 Hastings. 1882 E. obliqua. Ex 11-viii-1976 LH 9,12 soil.
8211-23 Hastings. 43 yr. E. obliqua. Ex 11-viii-1976 LH 5,9 soil.
8211-24 Stop 3 on Kermandie track from 24-viii-1973 JLH, AMMR 9 Hartz Mts. to Kermandie Plains. West of Taylor's Ridge. 8211-822149.
8211-25 On track from Hartz Mts. hut to 24-viii-1973 JLH, AMMR 4,9 Kermandie Plains. West of Tay- lor's Ridge. 8211-822149.
8211-26 On track from Hartz Mts. hut to 24-viii-1973 JLH, AMMR 9 Kermandie Plains. W of Taylor's Ridge. 8211-822149.
8211-27 Track from Hartz Mts. hut to 24-viii-1973 JLH, AMMR 5,6 Kermandie Plains. At head of Arve River, W of Taylor's Ridge. 8211-829146.
324
Sample No. Locality and Habitat Description Date Collector Species
8211-28 Track from Hartz Mts. hut to 24-viii-1973 JLH, AMMR 6 Kermandie Plains. At head of Arve River, W of Taylor's Ridge. 8211-829146.
8211-29 Northern junction of Arve Loop 2-iii-1976 LH 9 in Arve Valley. 8211-807273.
8211-30 On South Weld Rd., W from Tahune 29-v-1974 JLH et a/. 4,9,12 Bridge (92.8 miles). 8211-783309.
8211-31 North junction of Arve Loop, 2-iii-1976 LH 9,12 Arve Valley. 8211-807273.
8212-1 Burnt area, between two bridges 28-v-1975 JAF 12 over Tyenna River.
8212-2 Past National Park, beside road 28-v-1975 JAF 9,12 near old quarry site.
8212-3 Just off Port Davey track, 150 m 28-v-1975 JAF 12 from road. 8212-643646.
8212-4 In debris near Lake Dobson. 5 -ii -1962 9,12
8212-5 At beginning of climb to Lake 15-x-1978 JAF 4,9,12 Skinner. About 3/4 hour in from start of track.
8212-6 Just on E side of ridge at top 15-x-1978 JAF 4,12 of climb to Lake Skinner plat- eau. 8212-747455.
8212-7 Lake Skinner track. Just in 15-x-1978 JAF 4,9,12 rainforest on boundary of eucalypt woodland and rainforest.
8212-8 About 4 m 2/3 way back on track 15-x-1978 JAF 9,12 from Lake Skinner.
8212-9 Lake Dobson. Moss Ex Nothofagus. 24-ix-1977 PMcQ 12
8212-10 Lake Fenton. Mt. Field National 13-ii-1968 TM staff 8 Park.
8212-11 Lady Barron Falls. National Park 19-i-1928 GPW 9,12
8213-1 SW of Great Lake. 8213-836485 1-vi-1975 JAF 8,12
8213-2 On road between Lagoon of 1-vi-1975 JAF 12 Islands turnoff and Arthur's Lake. 8213-917406.
8214-1 On Quamby Bluff Road. 31-v-1975 JAF 12 8214-695000.
8214-2 Below Quamby Bluff. 8214-764895. 31-v-1975 JAF 12 325
Sample No. Locality and Habitat Description Date Collector Species
8214-3 Fairy Glade, on Quamby Bluff 1-vi-1975 JAF 12 Rd. Above road. 8214-767882.
8214-4 Just S of Pine Lake, at Pencil 1-vi-1975 JAF 12 Pine notice. 8214-755785.
8214-5 Golden Valley. Open dry sclero - 18-ix-1978 CR 12 phyll forest.
8215-1 Ex yabbie burrow at bridge over 20-vii-1974 AMMR et a/.12 Brown Creek. E side of Port Sorell. 8215-670424.
8215-2 Dazzler Range. SW side Ex leaf 21-vii-1974 JLH et a/. 12 litter.
8215-3 W slope of Holwell Gorge, at 20-vii-1974 JLH et a/.12,15 Picnic Area. Ex moss.
8215-4 E slope of Holwell Gorge, at 20-vii-1974 JLH et a/. 12 Picnic Area. Ex Casuarina litter.
8215-5 As for 8215-3, ex leaf litter. 20-vii-1974 JLH et al. 12,15
8215-6 As for 8215-4, ex leaf litter. 20-vii-1974 JLH et al. 12,15
8215-7 Green's Beach. In Acacia scrub 1-i-1980 JAF 13,15 just behind beach at W end. 8215-783518.
8311-1 SW of Pelverata, between road 27-v-1975 JAF 9 and Kellaway's creek. 8311-080328.
8311-2 Margate Beach, left of Beach 29-v-1975 JAF 12 Rd., in front of pines. 8311-224357.
8311-6 Beside Woodbridge Hill Rd. 29-v-1975 JAF 8,9,12 8311-165200.
8311-7 Adventure Bay, Forestry Rd. 29-v-1972 BK et al. 8,12 8311-258980.
8312-1 N edge of farm dam, E side of 7-ix-1972 BK 12 Tunnel Hill. 8312-343555.
8312-2 "Fern Tree Gully". 4-v-1972 8,9,12
8312-3 Beside a creek near Southern 27-v-1975 JAF 9,12 Outlet past Kingston. Small un- cleared patch on side of gully. 8312-190422.
8312-4 Past Mountain River - just near 27-v-1975 OAF 12 gate past sawmill. 8312-110477. 326
Sample No. Locality and Habitat Description Date Collector Species
8312-5 In gully at southern end of sand-14-iii-1972 BK 9 sandstone 4 miles S of Tunnack Rd. to Colebrook.
8312-6 Risdon, E side of Derwent River. 7-iii-1972 9 Ex grass tussock.
8312-7 Various habitats behind hut on 9-v-1972 JLH 8,9,12 Mt. Wellington at junction of Organ Pipes track and road. 8312-190512.
8312-8 Risdon. Ex grass tussock. 16-viii-1976 JLH, RBM 9
8312-9 682 Sandy Bay Rd. In a dry 13-xi-1977 JAF 9 drainage ditch in back garden.
8312-10 Royal Botanical Gardens, Hobart. 1-viii-1978 JAF 8
8312-11 Fern-Glade, Ferntree, Mt. Well- 2-ii-1979 JAF 8,12 ington.
8312-12 Browns Rd. area, unburnt. 2-ii-1979 JAF 8,9
8312-13 Royal Botanical Gardens, Hobart. 1-vii-1978 JAF 8
8312-14 Mt. Wellington pinnacle. 2-xii-1975 PMcQ 9
8312-15 Lambert Park Creek, on top side 19-ii-1979 JAS 8,9,12 of Churchill Ave., right bank.
8312-16 Taroona, in a suburban garden. 2-i-1978 HDB 8 TMAG G1938.
8312-17 Taroona. G1751. 30-xii-1975 JDS
8312-19 Ferntree, Mt. Wellington 19-ii-1928 8 SAM TC905.
8313-1 . NW corner of Lake Sorell, near 13-xi-1975 AMMR 12 lakeside. 8313-096379.
8313-2 Mountain Creek. Lake Sorell. 1-vi-1977 REM 12
8315-1 Parking spot going up Sideling 31-v-1975 JAF 12 from Scottsdale. 8315-365341.
8315-2 In small gully beside road near 31-v-1975 JAF 12 top of Sideling. 8315-358341.
8315-3 Beside creek at "Shingle Shed" 11-ix-1975 JAF 12 at end of road at APPM Phillip's block. 8315-308302.
8315-4 3 km SW of South Barrow. 840 m 12-xii-1978 CR 14 alt. 327
Sample No. Locality and Habitat Description Date Collector Species
8411-1 Eaglehawk Neck 14-v-1968 PSL 8
8411-2 At top of hill on NW-direction 20-iv-1977 AMMR, JAF, 8.9.12 section of end of Campbell's RGW Rd. 8411-645277.
8411-3 Campbell's Rd., near Koonya. 20-iv-1977 AMMR et a/.8,12 Just past big bend, on lower side of road. 8411-648273.
8411-4 At end of navigable part of 21-iv-1977 AMMR 5,12 Jetty Rd., on Eaglehawk Bay. 8411-732367.
8411-5 As for 8411-4, but lower, on 21-iv-1977 AMMR 5,8 upper beach. 8411-732367.
8411-6 First creek on Cape Raoul track. 31-xii-1976 JAF 8,9,12 8411-628165.
8411-7 1.5 km past lookout on track to 31-xii-1976 JAF 8,12 Cape Raoul. 8411-637148.
8411-8 S side of Eaglehawk Bay. Tasman 22-iv-1978 AMMR et al.7 Peninsula.
8411-9 Eaglehawk Neck, near turnoff 14-ii-1968 PSL 9 to Waterfall Bay.
8412-1 Pawleena Dam. 8-ix-1972 BK 8
8412-3 34.3 miles from Hobart on Tasman 15-viii-1972 JLH et a/. 9,12 Highway. Lower side of road.
8412-5 N end of Slopen Main Beach. Alt. 18-vi-1976 KF 8 30'. 8412-558417.
8412-6 Beside Prosser River, opposite 18-ix-1976 JAF 9 Burnt Bridge gully. 8412-681880.
8412-7 As for 8412-6 18-ix-1976 JAF 9
8412-8 Unemployed Gully, S of Btckland. 19-ix-1976 JAF 9,12 Just E of crossing, near entrance to gully beside Mt. Gatehouse. 8412-598787.
8412-9 Valley beside Mt. Gatehouse, 19-ix-1976 JAF 9,12 100 m from mouth. S of Buckland. 8412-599788.
8412-10 Just above E side of creek in 19-ix-1976 JAF 8,9,12 gully under Mt. Gatehouse, 130 in from junction of Unemployed Gully, S of Buckland. 8412-599788.
328
Sample No. Locality and Habitat Description :Date . Collector Species
8412-11 Top of Crawler's Gully, just 19-ix-1976 JAF 9,12 over creek flowing along bottom of Thunderbolt Hill. 8412-602675.
8412-12 Beside Mitchell's Creek on Buck- 9-ii-1977 JAF 9 land-Levendale Rd. 8412-564867.
8412-13 2 km S of Levendale, beside 9-ii-1977 JAF 12 creek, opposite pasture. 8412-465887.
8412-16 By Nelson Creek, on N side 19-xi-1975 JAF 8,9 above bridge. 8412-540821.
8412-17 Prosser Estuary. First gully up 2-i-1977 JAF 9,12,15 from bridge on N bank.
8413-1 Near TPFH road, in dense forest. 15-viii-1972 JLH et al. 8,9,12,15 8413-708195.
8413-2 Quarry site. E side of TPFH road 1-iii-1972 JLH, JM 8,9,12,15 in from Little Swanport. et al. 8413-697230.
8413-3 TPFH road, TO Collecting site. 1-vi-1972 BK 9,12,15 3 8413-698208.
8413-4 As for 8413-3. 1-vi-1972 BK 9,12
8413-5 Quarry site. S side of creek, 1-iii-1972 JLH et al. 8,9,12,15 W side of TPFH road in from Little Swanport. Ex litter. 8413-697230.
8413-6 Dense "rainforest" W side of 1-iii-1972 JLH 8,12,15 TPFH road from Little Swanport. Ex litter. 8413-698208.
8413-7 As for 8413-6. Ex moss. 1-iii-1972 JLH 12,15
8413-8 As for 8413-5. Ex moss. 1-iii-1972 JLH 12
8413-9 Woodchip road. East Coast. 1-iii-1976 ? 15 T08. Ex creek.
8413-10 Near gate, first bridge on A 18-ix-1976 JAF, JLF 9,15 road near Triabunna. 8413-150980.
8413-11 About half- way up hill on north- 18-ix-1976 JAF, JLF 9,12 ern side of 8413-10 valley near A road, near Triabunna. 8413-750982.
8413-12 Top of hill NW of first bridge 18-ix-1976 JAF, JLF on A road, near Triabunna. 8413-748984. 329
Sample No. Locality and Habitat Description Date Collector Species
8413-13 Crayfish Swamp, S of Lake Leake. 9-viii-1973 BK et al. 12,15 8413-705402.
8413-14 Beside Rocky Waterhole Creek. 19-xi-1975 JAF 8,9,12 Tasman Highway. 8413-768062.
8413-15 Creek before Percy's Gully on 19-xi-1975 JAF 12,15 M road. 8413-725155.
8413-16 Tiger Creek. E side of road. 22-xi-1975 JAF 12,15 8413-713183.
8413-17 As for 8413-16. 22-xi-1975 JAF 9,12,15
8413-18 Tiger Creek. W side of road. 22-xi-1975 JAF 12,15 8413-713183.
8413-19 As for 8413-18. 22-ix-1975 JAF 9,12,15
8413-20 Tower Road, off M road. 22-xi-1975 JAF 12,15
8413-21 As for 8413-20. 22-xi-1975 JAF 8,12
8413-22 Beside M road, in dry sclero- 22-xi-1975 JAF 9,12 phyll N of M16. 8413-707350.
8413-23 Mitchelmore's Creek on Swanston 19-xi-1977 JAF 9,12 Road. 8413-734119.
8413-24 Beside creek, N side of M28, off 9-viii-1973 JLH 9,12 M road. 8413-693422.
8413-25 S of "Windfalls", 12 miles SE of 30-v-1973 AJD 12 Campbell Town. TM G1635.
8414-1 Near hut at treeline on Ben 31-i-1971 JLH, BK 12 Lomond.
8415-2 Weld River, on banks. 31-x-1972 RBM, BK 12
8415-3 Myrtle forest (sign posted) just 30-v-1975 JAF 12 before bridge over Weld Fiver. 8415-780370.
8415-4 Near Mathinna Falls. 28-v-1973 RBM 12
8415-5 Mathinna Plains. Ex moss. 29-v-1973 JLH 12
8415-6 Mathinna Plains. Ex litter. 29-v-1973 JLH 12
8415-7 South Esk River at Mathinna. 28-v-1973 RBM 12
8415-8 Beside a tributary of S. Georges 9-xi-1977 AMMR et a/.12,15 R., near West Pyengana. 8415-776260.
8415-9 Beside Weld R., 4 km from 9-xi-1977 AMMR et a/.12 Moorina. 8415-735453. 330
Sample No. Locality and Habitat.Description. .Date Collector Species
8415-10 Between Russel's Rd. and Ben R 2-vii-1974 JMS, AS 12 Ridge Rd., off Maurice Rd. 8415-505210.
8415-11 Beside Simon's Rd., off Diddleum 2-vii-1974 JMS, AS 12 Rd. 8415-436219.
8415-12 One mile W of Trenah Bridge on 2-vii-1974 JMS, AS 12,15 Maurice Rd. 8415-558266.
8415-13 S of Rowlings Rd. and Red Hills 2-vii-1974 JMS, AS 12 Rd. junction, near Branxholm. 8415-578410.
8415-14 Between two easterly spur roads 2-vii-1974 JMS, AS 12 off Kamona Valley Rd. NW of Branx- holm. 8415-559455.
8415-15 At end of a westerly spur off 2-vii-1974 JMS, AS 12 Warrentinna Rd. from Branxholm. 8415-598443.
8415-16 Myrtle Creek. S of Mathinna. 15-vii-1973 JLH 12 Ex moss.
8415-17 St. Columba Falls area. Ex moss. 18 -vi -1974 PS 12,14,15
8415-18 10 miles from base ofSt. Col- Easter, 1978 BR 12,15 uMba Falls, near Pyengana. 8415-770250.
8415-19 As for 8415-17. Ex litter. 18-vi-1974 PS 12,15
8416-2 Near turnoff to Tomahawk. 31-v-1975 JAF 12 8416-651702.
8512-1 Maria Island. early 1977. MB et a/. 9,12
8512-2 As for 8512-1. early 1977 MB et a/. 9,12
8512-3 As for 8512-1. early 1977 MB et al. 9
8512-4 Maria Island. Open dry sclero- March, 1977 GE 9,12 phyll forest. 8512-888843.
8512-5 Maria Island. Above cliffs, 3-i-1977 JAF 9 where trees start on the way up Bishop and Clerk. 8512-888848.
8512-6 Maria Island. Near bottom of 3-i-1977 JAF 8,12 scree on W face of Bishop and Clerk. 8512-910841.
8513-1 Hepburn Point, Freycinet GE 8 Peninsula. 8513-037365. 33 1
Sample No. Locality and Habitat Description Date • Collector Species
8513-3 Schouten Island. In valley near 19-i-1978 JAF 8,12 the north coast, E of Crockett's Bay. 8513-075160.
8513-4 Schouten Island. E bank of creek 19-i-1978 JAF 12 flowing into Crockett's Bay (branch west of Bear Hill). 8513-058157.
8513-5 Schouten Island. Behind beach at 20-i-1978 JAF 8 Crockett's Bay. 8513-057159.
8513-6 Schouten Island. Dry bed of 20-i-1978 JAF 12 Chinese Creek, 100 metres below falls. 8513-052151.
8514-1 Bush beside road at Lower Ele- 30-v-1975 JAF 12 phant Pass. 8514-034874.
8514-2 N side of gully nr 8514-1 at 30-v-1975 JAF 12 Elephant Pass. 8514-034874.
8514-3 As for 8514-2, other side of 30-vii-1975 JAF 12,15 gully.
8514-4 Gully near bridge at top of 30-vii-1975 JAF 12,15 Elephant Pass near Lookout. E side of gully. 8514-034899.
8514-5 Top of Elephant Pass, other side 30-v-1975 JAF 12,15 of gully from 8514-4. 8514-034899.
8514-6 St. Mary's Pass. Ex litter. 30-v-1975 JAF 12,15 8514-026999.
8514-7 As for 8514-6, under sags. 30-v-1975 JAF 12,15
8514-8 Beside a tributary of the Break 10-xi-1977 AMMR et a/.15 O'Day River at Cornwall. 8514-946967.
8514-9 7 km E of St. Mary's. Elephant 21-x-1975 PMcQ 15 Pass.
8514-10 In Eucalypt forest 4 miles S of 13-viii-1974 PM 12 Gray. TM G1605.
8515-1 Beside Tin Dish Creek, near Lot- Feb. 1977 RBM 14 tah. 8515-842361.
8515-2 Great Musselroe River, 10.1 mile 1-xi-1972 RBM, BK 12,14 from Pioneer River turnoff. 8515-885545.
8515-3 Above Trafalgar Flat, on Hogan's 26-xii-1976 RBM 12 Road. 8515-863183. 332
Sample No. Locality and Habitat Description Date Collector Species
8515-4 Upper reaches of Scamander R., 26-xii-1976 RBM 12,15 on Hogan's Road. 8515-916163.
8515-5 "The Bottleneck", Ansons 8-xi-1977 AMMR et a/.12,14,15 River. 8515-034549.
8515-6 "The Bottleneck", Ansons R. 8-xi-1977 AMMR et a/.12,14 Small tributary of Ansons R., on right bank. 8515-034549.
8515-7 Old Chum Creek. 8515-894568. 7-xi-1977 AMMR et a/.12
8515-8 Peacock Creek. 8515-849602. 7-xi-1977 AMMR et a/.12,14
8515-9 Great Musselroe River, on 7-xi-1977 AMMR et a/.12,14 Tebrabuna Rd. 8515-892509.
8515-10 Last River - Ansons Bay Pd. 8-xi-1977 AMMR et a/.12,14 8515-006457.
8516-2 Brown's Bridge Rd., just W of 7-xi-1977 DC, AMMR 12 Brown's Bridge. 8516-912674. 333
Key to species code numbers.
Species Code
Orchestiella neambulans 1
0. guasimodo 2
Tasmanorchestia annulata 3
Neorchestia plicibrancha 4
Austrotroides longicornis 5
A. leptomerus 6
A. maritimus 7
Mysticotalitrus tasmaniae 8
M. crypticus 9
Arcitalitrus sp.S. 10
Keratroides albus 11
K. vulgaris 12
K. rex 13
K. pyrensis 14
K. angulosus 15 334
List of Collectors
Collector Abbreviation Collector Abbreviation
P. Allbrook PA P. McQuillam PMcQ
A.P. Andrews APA R. Mesibov PM
H.D. Barker HDB P. Nally PN
G. Bennison GB C. Oke CO
L. Bradshaw LB C.T. Peters CTP
M. Brett MB C. Reid CR
G. Burns GB A.M.M. Richardson AMMR
J. Burrell JB R. Russell RR
L. Chang LC B. Ryland BR
D. Coleman DC J.D. Skuja JDS
A.J. Dartnall AJD B.J. Smith BJS
G. Edgar GE J.A. Smith JAS
S. Evans SE A. Stonjak AS
K. Fagan KF J.M. Stonjak JINS
J. Fenton JF P. Suter PS
J.A. Friend JAF D. Swift DS
J.L. Friend JLF T.M. Walker TMW
B. Goede BG R.G. White RGW
J.J. Greenhill JJG G.P. Whitley GPW
V.V.Hickman VVH 7.L. k-vft&n. 71.164.
L. Hill LH
M. Hortle MH
J. King JK
B. Knott BK
P.S. Lake PSL
J. Madden JM
R.B. Mawbey RBM 335
APPENDIX II
Geographical locations and identifications of specimens of Talitridae found in fully terrestrial situations and reported in published work.
Specific and generic designations are as reported, irrespective of subsequent synonymy or re-allocation. References appear below the table, unless already listed in the main bibliography.
The world distribution of terrestrial Amphipoda shown in Figure 4.1 is plotted from these records, with the addition of localities in
Australia and New Guinea from which material has been found in
Australian museums. Other unpublished records mentioned in the text are also represented in Figure 4.1. 336
Author and Date Place Species recorded
Afonso, 1977 Azores Talitrus pacificus
Orchestia mateusi
Andersson, 1963 Canary Islands Talitrus alluaudi
Orchestia chevreuxi
0. platensis
Baker, 1915 Luzon, Philippines Parorchestia luzonensis
J.L. Barnard, 1955 Oahu, Hawaiian Is. Orchestia kaalensis
O. pickeringi
Parorchestia hawaiensis
J.L. Barnard, 1960 Micronesia:
Palau Is. Talitrus hortulanus
Orchestia floresiana
Yap Is. O. floresiana
Caroline Atolls O. floresiana
Truk Is. Talitrus hortulanus
T. toli
T. trukana
Ponape T. nesius
Orchestia ponapensis
Kusaie O. floresiana
Gilbert Is. O. floresiana
Fiji O. floresiana
O. floresiana vitilevana
K.H. Barnard, 1916 South Africa Talitriator eastwoodaet
K.H. Barnard, 1935 Burma Talitrus sp.
Cochin State, India Parorchestia notabilis
K.H. Barnard, 1936 Mauritius Talitroides topitotum
Orchestia mauritiensis 337
Author and Date Place Species recorded
K.H. Barnard, 1940 South Africa Talitroides eastwoodae
forma typica
forma cylindripes
forma setosa
forma calva
forma macronyx
K.H. Barnard, 1958 Madagascar Talitroides topitotum
Bate, 1862 New Zealand Orchestia telluris
0. sylvi cola
Biernbaum, 1980 S. Carolina Talitrus alluaudi
T. topitotum
Bousfield, 1964 Campbell I. Parorchestia campbelliana
P. insularis
Bousfield, 1971 Tahiti Orchestia floresiana
New Ireland O. anomala
New Britain O. sp.
Hermit Is. Parorchestia macrochela
P. similis
Mussau Brevitalitrus wolffi
Dyaul B. dyaulanus
Bousfield, 1976 Lord Howe I. Parorchestia gowerensis
Bousfield & Carlton,
1967 California Talitrus sylvaticus
Bousfield & Howarth,
1976 Hawaiian Is. Spelaeorchestia koloana
Talitroides alluaudi
T. topitotum
Bowman, 1977 Galapagos Is. Orchestia vaggala
Burt, 1934 Ceylon Talitrus (Talitropsis) topitotum 338
Author and Date Place Species recorded
de Castro, 1972 Brazil Talitrus pacificus
Carl, 1934 Southern India Talitrus decoratus
Chevreux, 1888 Canary Is. Orchestia chevreuxi
Chevreux, 1889 Azores 0. guernei
Chevreux, 1896 Seychelles Talitrus alluaudi
Chevreux, 1900 Canary Is. Orchestia chevreuxi
Azores O. guernei
Chevreux, 1901 Seychelles Talitrus alluaudi
Madagascar T. alluaudi
Chevreux, 1907 Gambier Archipelago T. alluaudi
Tahiti Talorchestia rectimana
Chevreux, 1915 New Caledonia T. antennulata
Parorchestia sarasini
Loyalty Is. Talorchestia antennulata
Chilton, 1911 Auckland Is. Parorchestia maynei
P. parva
Campbell I. P. insularis
Snares Is. P. improvisa
Stewart I. P. improvisa
Chilton, 1911 Kermadec Is. P. sylvicola
Chilton, 1912 Java Orchestia parvispinosa
Chilton, 1917 New South Wales Talitrus sylvaticus
Chilton, 1921 Juan Fernandez Is. Orchestia chiliensis
var.gracilis
Chilton, 1923 New South Wales Talitrus sylvaticus
South I. New Zealand Parorchestia sylvicola
Chilton, 1925 Chatham Is. P. sylvicola
Dahl, 1950 Canary Is. Orchestia canariensis
O. chevreuxi 339
Author and Date Place Species recorded
Dahl, 1950 (cont'd) Madeira Orchestia platensis
Dahl, 1967 Azores 0. chevreuxi
Talitrus pacificus
T. alluaudi
Madeira T. pacificus
Dana, 1853 & 1855 North I., New
Zealand Orchestia sylvicola
Tahiti O. rectimana
Hawaiian Is. O. hawaiensis
O. pickeringii
De Sylva, 1959 Ceylon Talitrus (Talitropsis) fernandoi
Duncan, 1968 South I., New Zealand Orchestia hurleyi
Duncan, 1969 South I., New Zealand O. hurleyi
O. patersoni
Friend, 1979 Tasmania Talitrus (Keratroides) vulgaris
T. (Keratroides) angulosus
Guerne, 1887 Canary Is. Orchestia chevreuxi
Hale, 1929 South Australia Talitrus kershawi
T. sylvaticus
Haswell, 1880 New South Wales T. sylvaticus
Haswell, 1881 Tasmania T. assimilis
Hunt, 1925 Scilly Isles T. dorrieni
Tasmania Parorchestia ? sp.
MIT-ley, 1955 Central Ck.,
Australia Talitrus sp.
New South Wales T. pacificus
Norfolk I. T. pacificus
Tasmania T. kershawi
North I., New Zealand T. sylvaticus
South I., New Zealand T. sylvaticus 340
Author and Date Place Species recorded
Hurley, 1955 (cont'd) Australia Talitrus sylvaticus
Hurley, 1957 North I., New Zealand Orchestia tenuis
0. lesliensis
O. rubroannulata
South I., New Zealand O. tenuis
O. lesliensis
O. sinbadensis
Snares Is. O. improvisa
O. patersoni
O. simularis
Stewart I. O. patersoni
Bench I. O. patersoni
Solander I. O. patersoni
Karewa I. O. rubroannulata
Little Barrier I. O. rubroannulata
Stephens I. O. rubroannulata
Kapiti I. O. rubroannulata
Auckland Is. O. parva
O. maynei
Campbell I. O. insularis
Hurley, 1968 Jamaica Not identified
Haiti' 11 11
Barro Colorado I. 11 11
Ingle, 1958 Cornwall, U.K. Talitrus sylvaticus
Iwasa, 1939 Hokkaido Orchestia platensis japonica
O. kokuboi
O. solifuga
Honshu O. platensis japonica
Danzyo I., Japan O. platensis japonica 341
Author and Date Place Species recorded von Martens, 1868 Japan 0. humicola
Mallis, 1942 California Talitrus sylvaticus
Miers, 1876 Rodriguez T. gulliveri
Monod, 1970 Galapagos Is. Orchestia costaricana
Monod, 1975 Reunion Talitrus sp.
T. topitotum
Mauritius T. gulliveri (?)
Murphy, 1973 Cornwall, U.K. T. dorrieni
Murphy, 1974 Cornwall, U.K. T. dorrieni
Rawlinson, 1937 Ireland T. dorrieni
Ruffo, 1947 Annobon I. T. gulliveri
Ruffo, 1949 New South Wales T. sylvaticus
Tasmania T. sylvaticus
T. tasmaniae
New Guinea Orchestia kinabaluensis (?)
Ruffo, 1958 Comoro Is. Talitrus pacificus
Reunion T. pacificus
Madagascar T. pacificus
Ruffo & Paiotta, 1972 New Caledonia T. sp.
Sandell, 1977 Victoria T. kershawi
T. sylvaticus
Sayce, 1909 Victoria T. sylvaticus
T. kershawl
Schellenberg, 1934 New South Wales T. dorrieni
Schellenberg, 1938 Bismarck Arch. Orchestia anomala
Shoemaker, 1935 North Borneo Parorchestia kinabaluensis
Shoemaker, 1936 California Talitrus sylvaticus
Louisiana T. sylvaticus
Shoemaker, 1942 Clipperton I. Orchestia marquesana 342
Author and Date Place Species recorded
Stebbing, 1900 Hawaiian Is. Orchestia platensis
0. pickeringii
Parorchestia hawaiensis
Stebbing, 1906 New Zealand P. tenuis
Stebbing, 1917 South Africa Talitriator africanus
Stebbing, 1922 South Africa Talorchestia sp.
Stephensen, 1927 Auckland Is. Parorchestia maynei
P. insularis
P. insularis var.
P. parva
P. sp.
Stephensen, 1935a Tahiti Talorchestia rectimana
Moorea T. rectimana
Stephensen, 1935b Marquesas Talitrus sylvaticus
Orchestia floresiana
O. floresiana f. monospina
O. marguesana
Hawaiian Is. Talitrus sylvaticus
New Britain Orchestia floresiana
Stephensen, 1938 Stewart I. Talorchestia patersoni
Parorchestia improvisa
P. stewarti f, brevicornis
f. longicornis
Campbell I. P. insularis
P. sp.
Stephensen, 1943 Samoa Talitrus hortulanus
New Hebrides T. hortulanus
Scilly Isles T. sylvaticus
Smith, 1909 Tasmania T. sylvaticus 343
Author and Date Place Species recorded
Tattersall, 1914 Assam Talorchestia kempi
Tattersall, 1922 Singapore T. malayensis
Tattersall, 1929 New Hebrides Orchestia platensis
Tamura & Koseki, 1974 Japan Orchestia platensis japonica
Thomson, 1881 South I., New Zealand 0. sylvicola
Campbell I. O. sylvicola
Stewart I. O. sylvicola
Thomson, 1892 Tasmania Talitrus sylvaticus
Deno, 1929 Japan Orchestia kokuboi
Watanabe, 1970 Japan O. platensis japonica
Weber, 1892 Flores O. floresiana
Java O. pervispinosa
Celebes O. montane 344
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These articles have been removed for copyright or proprietary reasons.