Darter Reproductive Seasons Author(s): Clark Hubbs Reviewed work(s): Source: Copeia, Vol. 1985, No. 1 (Feb. 11, 1985), pp. 56-68 Published by: American Society of Ichthyologists and Herpetologists (ASIH) Stable URL: http://www.jstor.org/stable/1444790 . Accessed: 10/01/2012 14:26

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http://www.jstor.org 56 COPEIA, 1985, NO. 1

changes in kinosternid turtles. J. Herpetol. 6:183- . 1938. Seasonal changes in the testes of the 189. musk turtle Sternotherusodoratus L. J. Morphol. 63: MCPHERSON, R. J., AND K. R. MARION. 1981. Sea- 301-317. sonal testicular cycle of the stinkpot turtle (Ster- SAINTGIRONS, H. 1982. Reproductive cycles of male notherus odoratus) in central Alabama. Herpetolog- snakes and their relationships with climate and fe- ica 37:33-40. male reproductive cycles. Herpetologica 38:5-16. MITCHELL, J. C. 1982. Population ecology and de- SPEAT, R. H. 1973. Seasonal variation in the tubular mography of the freshwater turtles Chrysemyspicta and interstitial areas of the testes in Sternothaerus and Sternotherusodoratus. Unpubl. PhD Dissertation odoratus L. Unpubl. MS Thesis, East. Ill. Univ., Univ. , Knoxville. Charleston. . (in press). Variation in the male reproductive STEEL,R. G. D., ANDJ. H. TORRIE. 1960. Principles cycle in a population of painted turtles, Chrysemys and procedures of statistics. McGraw-Hill Book Co., picta, from Virginia. Herpetologica. New York. MOLL, E. 0. 1973. Latitudinal and intersubspecific TINKLE, D. W. 1961. Geographic variation in repro- variation in reproduction of the painted turtle, duction, size, sex ratio and maturity of Sternotherus Chrysemyspicta. Herpetologica 29:307-318. odoratus(Testudinata: Chelydridae). Ecology 42:68- . 1979. Reproductive cycles and adaptations, 76. p. 305-331. In: Turtles perspectives and research. ZAR,J. H. 1974. Biostatistical analysis. Prentice-Hall, M. Harless and H. Morelock (eds.). John Wiley and Inc., Englewood Cliffs, New Jersey. Sons, New York. ZUG, G. R. 1972. Walk pattern analysis of crypto- NIE, N. H., C. H. HULL, J. G. JENKINS, K. STEIN- diran turtle gaits. Anim. Behav. 20:439-443. BRENNERAND D. H. BENT. 1975. Statistical package for the social sciences. McGraw-Hill Book Co., New DEPARTMENT OF BIOLOGY, UNIVERSITY OF York. RICHMOND, RICHMOND, VIRGINIA 23173. Ac- RISLEY,P. L. 1933. Observations on the natural his- cepted 20 April 1984. tory of the common musk turtle, Sternotherusodor- atus (Latreille). Pap. Mich. Acad. Sci. Arts Lett. 17: 685-711.

Copeia, 1985(1), pp. 56-68

Darter Reproductive Seasons

CLARK HUBBS

Twenty-nine darters have reproductive intervals demonstrated and geographic variation is shown for three species based on ten or more populations. The data from most populations are in accord with an hypothesis of multiple spawning by individual females. The length of darter reproductive seasons varies with latitude. Populations from the southern United States breed over longer intervals than those from more northern locations. Most of the difference is based on earlier initiation in southern than in northern regions; whereas, the end of the breeding season tends to be similar at different latitudes. The spawning season is longer in stenothermal habitats than it is in eurythermal habitats (of various latitudes). The data are in accord with hypotheses that photoperiod plays a major role in reproductive initiation and temperature is important in reproductive termination.

REPRODUCTIVE seasons have been ascer- peated efforts. Although Page (1983) opined tained for many darters (Page, 1983; that only one clutch was produced each season, Kuehne and Barbour, 1983); however, most re- my data suggest long reproductive seasons with relate ports to observations of spawning occur- multiple spawnings (Hubbs, 1983). Addition- rence during collection intervals with few re- ally, Gale and Deutsch (1985) provide data dem-

? 1985 by the American Society of Ichthyologists and Herpetologists HUBBS-DARTER REPRODUCTIVE SEASONS 57 onstrating multiple spawnings for females; it is presumed that females in natural olmstedi in Pennsylvania. My opinion that most populations were equally ripe at additional in- darters have multiple clutches has been ex- termediate dates as all samples from within the pressed in several papers but the data base has appropriate reproductive seasons had numer- been presented in detail only in Hubbs and ous reproductive females. The data reported Strawn (1957a) who showed that groups of ova are for occurrence of reproductive females, thus sequentially differentiated from the pool of im- the seasons recorded may be shorter than those mature ova. Herein, I report known dates of that occur in nature as some females may have reproduction for several populations of three been reproductive before and/or after those darters and for one or more populations of 26 dates recorded in my data. additional species. Those data are in general The localities reported here are the same as agreement with "Natural History" data pre- those reported in Hubbs (1967). Commonly, sented by Page (1983) and Kuehne and Barbour adjacent localities are pooled. Most sampling (1983) except that my information often sub- was from the years 1954 through 1967. As sev- stantially extends the previously reported re- eral leap years occurred within that time frame, productive seasons. Feb. is considered to have 29 days.

MATERIALS AND METHODS ETHEOSTOMA LEPIDUM

The data reported here are derived primarily The greenthroat darter reproductive season from my previous hybridization experiments has been reported as Oct. or Nov. through May (Hubbs, 1967). The procedures were to obtain with populations in stenothermal environments darters from a variety of geographic locations having a longer spawning season than those in and return them to Austin. The stocks were more eurythermal environments (Hubbs and maintained without feeding at ca 15 C. Females Strawn, 1957a). These data reported here (Fig. with clear, translucent eggs were considered to 1) corroborate that report. It should be noted be ripe and used for hybridization (or control) that the more stenothermal environments (S. experiments. During most experiments some Concho, Mt. Home, S. Guadalupe) have longer eggs developed embryos that hatched into suc- recorded reproductive seasons than the more cessful larvae; therefore, this conservative mea- eurythermal extensively collected environ- sure of reproduction is used here. Only those ments (Hunt, Ingram, Junction). The green- females whose eggs were fertilized are reported. throat darter is basically a springrun species. If I were to use clear, translucent eggs as the Thus, it is scarce or absent from very eury- index for reproduction, an occasional addition- thermal locations. Collectively (all populations al reproductive day would have been reported pooled) greenthroat darter eggs are shown to (especially for species that were tested infre- have been fertilized on 178 of 181 days in Nov. quently and their reproductive criteria not as through April (the exceptions are 20 Feb., 31 well understood). Females held under those lab- March, 25 April). Thus the species is intensively oratory conditions ceased producing eggs in 4 reproductive for six months and reproduction to 5 days after 50 to 90% of them had ovulated. is also recorded for Sept., Oct., May and June, Consequently, it is apparent that at least half, or at least some level of reproduction demon- and presumably all, females were in reproduc- strated for 10 months. Additionally, I record tive cycle at the times of capture. Samples early translucent eggs from females obtained in Blue or late in the reproductive seasons reported here Spring, , in Aug. 1983; Hubbs and or of poorly understood species may have had Strawn (1957a) recorded breeding females smaller fractions of females reproductive, but (based on translucent eggs) in July. this occurred in fewer than 1% of the samples. At the South Guadalupe locality, greenthroat For the data reported here female capture was darter females were reported to be reproduc- up to five days prior to the recorded date. Each tive in June, July and Aug. (Hubbs and Strawn, date is considered as a datum even though that 1957a) (the present report records fertilized eggs date may have been represented by 50 ripe fe- from that locality in the other nine months). males on each of three separate years. Each year That paper also reported spawning (the number had similar reproductive seasons, thus pooling of eggs spawned was equivalent to the number of years did not distort the data base. of ripe eggs in dissected females) in the labo- The data are for known occurrence of ripe ratory (with densities equivalent to those on 58 COPEIA, 1985, NO. 1

Etheosfoma lepidum

I I I I I I I I I

S. Concho- emoemoem. ? - - " - * - San Saba- Junction- Austin- Kerrville- Mt.Home- N. Guad - Ingram- Sept Oct Nov Dec Jon Feb Mar Apr May Jun Hunt- S. Guod- Frio - Fig. 1. Known reproductive dates for Etheostoma lepdum from 12 regons. Nueces-

Fig ] Known. . reproductivedates for Etheostomalepidum from ]2 regions.

spawning riffles) every 4.9 days at 19.6 C and are seldom collected and thus seldom recorded every 4.8 days at 20.4 C [the average temper- as ripe. The data reported here parallel those ature of central springs is ca 20 C (Hubbs, previously reported for the same species in Tex- 1971)]. As Hubbs and Martin (1965) showed as by Hubbs (1961) and by Hubbs et al. (1968). that greenthroat darter females lay eggs pri- They do extend the initiation of reproduction marily in daylight, it is assumed that the clutches into late Oct. Marsh (1980) has shown that the in nature occur every five days (previous aver- termination of spawning may be delayed until ages rounded to 5 days as nocturnal spawnings July in artificially cool hypolimnetic release unlikely). A female from the South Guadalupe waters in the Guadalupe River downstream from would lay 52 clutches per year based on the Canyon Reservoir. The natural reproductive known interval of fertilized eggs or 73 clutches season in Texas would thus be seven months based on observations of translucent eggs. The with a seminatural nine-month season. Assum- egg volume approximates 7% of female volume. ing a five-day interbrood interval this would If we use a conservative figure of 5% of female mean 42 to 54 spawns per year. volume, the summed annual contributions are The reproductive season in extends 260 and 365% of female volume. from mid-Feb. to early June, and in from early April to early June (Fig. 2). This ETHEOSTOMA SPECTABILE latter period is in accord with a March through May reproductive season in (Cross, 1967) The orangethroat darter has a recorded re- and Missouri (Pflieger, 1975) but suggests my productive season in central Texas that is very first Missouri samples were well after the first similar to that for the greenthroat darter. The spawning. The Arkansas season corresponds major difference is that orangethroat darters with or is slightly longer than that of late Feb. occupy more eurythermal environments, thus or March to May reported by Miller and Ro- lack the year-round reproduction that prevails bison (1973) for . The Arkansas pat- for greenthroats occupying springruns (Fig. 2). tern occurs in both the White River (E. s. spect- For example at Junction both species are first abile) and River (E. s. pulchellum) recorded as reproductive on 12 Nov. In gen- tributaries (contrast White R. and Clear Cr.). eral, reproduction is recorded over longer in- The spawning season of orangethroat darters tervals for the stations at which that species is varies with latitude. In the south, it extends for more abundant (North Guadalupe, Hunt, Kerr- seven months, farther north (Arkansas) it lasts ville, Austin for orangethroats) (S. Concho, In- almost four months, yet farther north (Mis- gram for greenthroats). Relatively rare species souri) it lasts three months and toward the -

HUBBS-DARTER REPRODUCTIVE SEASONS 59

Etheosfoma spectobile I I I I I I I I I 1 1 1II I I r Loose Cr. - Mo. I Maries - [ White R. - w ?~~~~?+ e? _1 - elele ?elll ? l ! Brush Cr. - Ark- LittleWildcat - -~~~ I SP~~ a ~~~~~~~lm eel ~ ~ ~~elOM Clear Cr. - . e e l * ? ? ? ee Mt, Cr. - Oklahoma - Lampassas - Solado - so 0 ?0 - ali elmOemei --::_- : eeelM Georgetown - MO? ellSM -el *W Me *? 4? e * 6 * OW *0 OM *o e * QM WW o * es *eo # * * t S. Concho - ~- e~ ~~~~ee *lee *ltl e ?* *llll ve vli e* *l ? e Junction - Austin -

*" -0 *-0 go 40 0 00 Me a el ? San Marcos - -@ # so e,,.-I4W 04 ....*:llelllelam ee*le0 a leil!i": "- *" :: ii Comfort- - *- . ~alIitI ~ I I ~t 1411 ~* / *I _1??l i I I I ,Ite iii i e l I* eeelI** I* ee iiI* ii e i Kerrville - I C ?I t I i i i itielII I II tI N. Guad.- Ingram - -r emW* -e e-mme -*? * mo t * -- ee *.1 i Hunt - el ? ell NlMIli eeleHii? * 4il* e? ll I I i I t I I Oct Nov Dec Jan Feb Mar Apr May Fig. 2. Known reproductivedates for Etheostomaspectabile from 20 regions. northern limit (Michigan) two months [early bour (1983). My Texas stocks would be P. car- April to early June (Winn, 1958)]. Winn's re- bonaria and my Oklahoma, Arkansas and Mis- port of peak reproduction in Kentucky and souri stocks P. caprodes if the two taxa are Tennessee during the first week in April cor- considered distinct species. responds better with the Missouri-Kansas sea- The Texas stocks have been recorded as son than with that of Michigan. The majority breeding fromJan. throughJune (Hubbs, 1961) of the north-south difference in breeding sea- and Oct. and Nov. (Stevenson, 1971). The latter son lies in the initiation [Oct. (Texas) to April report relates to spawning migrations and pre- (Michigan)] rather than the end [mid-May (Tex- sumably involves a prespawning migration and as) to early June (Arkansas-Missouri-Michi- not actual spawning. The locality referenced by gan)]. Assuming equivalent interbrood intervals Stevenson is in the Kerrville locality included and equivalent individual clutch numbers in in Fig. 3. That locality has a recorded breeding equal-sized females, Michigan females would season of Jan. through May (presumably also have to survive 3.5 spawning seasons to produce late Dec.) but definitely not including Nov. dur- the same number of offspring as Texas females ing which month many were exam- would produce in one year. ined at the same time that orangethroat darter females are reproductive. The present data CAPRODES demonstrate that the season is mid-Dec. to mid- May. The logperch has been considered to repre- The logperch is commonly found in the same sent two species (the southern P. carbonaria and streams as the orangethroat darter. The two the northern P. caprodes)by Thompson (1978), species tend to partition riffles with logperch in an action endorsed by Morris and Page (1981), the deeper water or associated with larger rocks and Page (1983) but not by Kuehne and Bar- than the orangethroat darter. Nevertheless the 60 COPEIA, 1985, NO. 1

Percina coprodes

II I I I I I I I I Missouri - Arkansas- Oklahomo- Brozos Syst.- ?? San Saba R.- ?? ? ? ?? Junction- ? ? ?Y ?? ?? Pedernales R.- ? Y Austin- ? ??? Cummins Cr - San Marcos- ?? ??? Gonzales- Comfort- ??? ?? ?? Kerrville- ??? Ingram- ? ? Hunt-

Dec Jan Feb Mar Apr May Fig. 3. Known reproductivedates for Percinacaprodes from 15 regions.

two species are commonly taken in the same March or April for Oklahoma); Missouri, April seine haul. Therefore, logperch females were and May (Pflieger, 1975) (my data cover only available for examination for ripe eggs on most May); southern Michigan, April to late May or of those dates orangethroat darters were ex- early June, and northern Michigan, mid-June amined. Both species had numerous reproduc- to late July (Winn, 1958). Except for northern tive individuals at five localities (Ingram, Kerr- Michigan (where orangethroat darters do not ville, Comfort, Austin,Junction). At each locality occur) the pattern is very similar to that of or- orangethroat darters were reproductive before angethroat darters, progressively shorter logperch (+56, +52, +24, +41 and +52 days, spawning seasons with more northerly latitudes. respectively, x = 45 days earlier), but the data The majority of the change is in spawning ini- on last reproduction are more equivocal (-3, tiation and little difference in the date of the +7, -29, -14, 0, x = 8 days later). The obvious end of the reproductive season. conclusion is that the breeding season for log- starts later but ends at about the same MISCELLANEOUS PERCINA time as that for orangethroat darters in central Texas. That season would be mid-Dec. or early Bigscale logperch.-The reproductive season for Jan. to mid-May. In a similar fashion, the re- Percina macrolepida in its natural range has not productive season for logperch in Arkansas and been previously recorded. Ripe females are Missouri starts later but ends at about the same known to occur from 26 Feb. to 14 April [all date as that of orangethroat darters. from central Texas localities (Fig. 4)]. The oc- The breeding season for male logperch is sub- currence of small individuals (ca 25 mm SL) in stantially longer than that for female logperch Lake Texoma in early June suggests a similar as various Etheostoma x P. caprodeshybrids were reproductive interval there. The available data produced in Nov., a month before the initiation on male reproduction show no additional re- of female reproduction. This observation is in productive interval. accordance with the generalization that for most fishes males are reproductive earlier (and long- Dusky darter.-The breeding season for Percina er) than females (Lagler et al., 1977; Nikolski, sciera has been reported as late May to earlyJuly 1963). in Illinois (Page and Smith, 1970) and Feb. to The spawning season for female logperch in June in Texas (Hubbs, 1961). My present data Texas is Dec. to mid-May; Arkansas, mid-March slightly expand that reported two decades ago to mid-May (Miller and Robison, 1973, list late with substantialJan. reproduction recorded. In HUBBS-DARTER REPRODUCTIVE SEASONS 61

Percina and Ammocrypta Species iI I I It ti I A. vvox E. Texas- P evides Arkonsas- P phoxocephola Missouri- P copelondi Oklahoma- P nigrofascialo - P shumardi Gonzales- P sciero E. Texas- Austin- San Marcos- *am- e "am1 0 ~o1 Oa ? Gonzales- *? ?* la ?o*r m ?* ee ** P mocrolepido C.Texas-

I I I I I I Dec Jan Feb Mar Apr May Jun Fig. 4. Known reproductivedates for seven species of Percinaand Ammocryptavivax. the reasonably stenothermal waters of the San Manitoba (Scott and Crossman, 1973). Starnes Marcos River, dusky darters are known to be (1977) opines that in Tennessee the river darter reproductive from 12 Jan. to 9 June (Fig. 4). spawns in Feb. and March. My data apply only Other localities (with more thermal variation) to one location in the Guadalupe River near have not been shown to have reproductive in- Gonzales, Texas, where river darters are shown dividuals for such a long interval but for the to be reproductive from 14 Jan. to 22 April, or most part have not been sampled as frequently. for more than 3 months (Fig. 4). The evidence The reproductive initiation for male dusky on male reproductive intervals is essentially the darters is considerably earlier than for the fe- same as that for females. males recorded here. Several males from Austin The recorded reproductive season for river had visible sperm that fertilized eggs (of Ethe- darters in Texas is longer than that for any ostoma)in late Dec. and early Jan. The first date other geographic area. Reproductive initiation was 29 Dec., or nearly a month before any fe- is markedly earlier in the south and the end is males from that population have been shown to somewhat later in the north. be ripe. The reproductive interval of dusky darters in Blackbanded darter.-The reproductive season San Marcos is five months, but in Illinois Page for Percina nigrofasciatahas been reported to be and Smith (1970) list an interval of less than two Feb. to April in Louisiana and early May to early months. Populations from all four Texas areas June near Auburn, Alabama (Mathur, 1973). have known reproductive intervals of three or My record from eastern Louisiana on 29 April more months. (Fig. 4) suggests that all of April is within the reproductive interval in Louisiana. River darter.-The reproductive season for Per- cina shumardi has been reported to be April in Channel darter.-The reproductive season for Kansas (Cross, 1967), April and May in Illinois Percina copelandi has been reported as 9-23 July (Thomas, 1970) and possibly June or July in in northern Michigan (Winn, 1958) and April 62 COPEIA, 1985, NO. 1

Texas Etheostoma

E. fonticolo Oct Nov Dec Jon Feb Mor Apr May Jun Jul

E. histrio E.Texos- E. osprigene E.Texas- E chlorosomum E.Texas- E. grocile E.Texas- E. proeliore E.Texos- E. rodiosum Okla.- E.Texas- E. grohomi Rio Gonde System Nov Dec Jan Feb Mar Apr May Fig. 5. Known reproductive dates for eight species of Etheostomafrom Texas and Oklahoma. Note the data for Etheostomafonticola are on a different scale.

and May in Kansas (Cross, 1967). The known scaly sand darter. Williams (1975) noted that reproductive season in Oklahoma extends from tuberculate males occur from mid-April to mid- 4 April to 29 June (Fig. 4). The two June data Aug. My record of a ripe female in east Texas points were obtained by Hubbs and Bryan (1975) on 8 April (Fig. 4) is the only report on a female from stocks obtained from the Blue River near reproductive date. I also record fertilization of its spring source (a stenothermal location). The darter eggs by a male scaly sand darter on 5 breeding season in the north is substantially April. shorter than that in the south.

TEXAS/OKLAHOMA ETHEOSTOMA Slenderhead darter.-The reproductive season for Percina phoxocephala has been reported to darter.-The reproductive interval be late May to early July in Illinois (Page and of Etheostomagrahami has been recorded as late Smith, 1971) and late April into May in Missouri March to early June (Harrell, 1980) but Strawn (Pflieger, 1975). My data for a Missouri stock and Hubbs (1956) report that spawning will oc- are supportive of Pflieger's report (Fig. 4). cur at regular intervals if the temperature re- mains near 20 C. My record of early Jan. ripe Gilt darter.-The reproductive season for Per- females (Fig. 5) is in accord with a prolonged cina evides has been reported to be in May in reproductive interval in stenothermal waters. It Virginia (Denoncourt, 1969). Pflieger (1975) is likely that the reproductive cycle for the Rio observed breeding males in late May in Mis- Grande darter is essentially the same as that of souri. I have obtained ripe females in late April the greenthroat darter. and early May in Arkansas (Fig. 4). Orangebellydarter.-The reproductive interval AMMOCRYPTA for Etheostomaradiosum has been reported to be March through May for the Blue River, Okla- My data relate to only one species of sand homa, by Scalet (1973). My data for Oklahoma darter, Ammocryptavivax. There is essentially no populations are in accord with Scalet's report information on the reproductive season of the but in east Texas reproduction (classification HUBBS-DARTER REPRODUCTIVE SEASONS 63 based on Hubbs, 1982) extends from at least Arkansas/Missouri - Etheostoma late Feb. through mid-April (Fig. 5). I I I I E. caeruleum Cypressdarter.-The reproductive interval for Etheostoma has been to be Mo.- proeliare reported April Ark.- and May in Illinois and as early as Jan. in Lou- E. isiana (Burr and Page, 1978). My data suggest nianguae prolonged spawning from earlyJan. to mid-April Mo.- in east Texas (Fig. 5). E julioe Ark.- Slough darter.-The reproductive season of E. terozonum Etheostomagracile has been reported to be March Mo.- in Texas (Collette, 1962) and late May and early E euzonum in Illinois and Smith, June (Braasch 1967). My Ark.- data indicate the breeding season starts in Tex- E. stigmaeum as (east) at least by 24 Jan. (Fig. 5). The dem- Ark.- onstrated Texas reproductive season is more than two months and that reported for Illinois E flabellore is no more than a month. Mo.- E. blennioides Bluntnose darter.-The reproductive season of Mo.- Etheostomachlorosomum has been reported to be Ark.- April in Kansas (Cross, 1967) and May in Illinois E. punclulatum (Smith, 1979). My data for Texas populations Ark.- demonstrate to late March early Jan. reproduc- E. zonale tion in east Texas The known (Fig. 5). spawning Mo.- season in Texas populations is longer than that for more northern localities. Ark.- reported I! I I I I I Feb Mar Apr May Jun Mud darter.-Little is known of the reproduc- 6. Known tive season of Etheostomaasprigene. Forbes and Fig. reproductivedates for ten species of Etheostomafrom Arkansasand Missouri. Richardson (1908) suggested a March to May reproductive season for Illinois and Becker (1983) stated that spawning occurs in April and ARKANSAS/MISSOURI ETHEOSTOMA May at Havana, Illinois. In east Texas repro- ductive females are recorded from 14 Feb. to Banded darter.-The reproductive season for 26 March (Fig. 5). Etheostomazonale has been reported as April and May in Oklahoma (Miller and Robison, 1973) Harlequin darter.-Little is known of the repro- and Kansas (Cross, 1967), April in Missouri ductive season of Etheostomahistrio. Kuehne and (Pflieger, 1975) and May and June in Illinois Barbour (1983) reported females apparently (Forbes and Richardson, 1908) and Pennsyl- to in ready spawn mid-March in and vania (Lachner et al., 1950). Additionally, Burr Hubbs and Pigg (1972) reported ripe females and Mayden (1979) record ripe females on 20 in Feb. That record is the 13 Feb. spawning July in western Kentucky. My data show Ar- recorded here Additional (Fig. 5). east Texas kansas reproduction from late March to early is recorded for 28 and spawning 29 March. June and Missouri reproduction from mid-April to late May (Fig. 6). The longest recorded re- Fountain darter.-Etheostoma fonticola has been productive interval is for Arkansas stocks recorded to be reproductive in all months in (southern). San Marcos River (Strawn, 1956). Schenck and Whitesides (1977) reported apparent spawning Stippled darter.-The reproductive season for in and late winter. peaks Aug. My data (Fig. 5) Etheostomapunctulatum has been reported as May fertilized documenting eggs during 10 months (Miller and Robison, 1973), April and May with are and (the exceptions Aug. Sept.) support males in breeding color as late as July (Pflieger, Strawn's thesis of continuous spawning. 1975), May (Kuehne and Barbour, 1983) and 64 COPEIA, 1985, NO. 1 probably April (Cross, 1967). My data, primar- as April and May (Pflieger, 1975). My data are ily based on a stock in a spring near Fayetteville, in accord with that season with ripe females Arkansas, have the breeding season at least from occurring between 2 April and 30 May (Fig. 6). mid-Feb. through mid-May (Fig. 6). As this species has a relatively restricted geographic Yokedarter.-The reproductive season for Ethe- range (Cloutman, 1980), it is likely that my data ostomajuliae has been reported to be in May in would apply to stenothermal waters (a preferred Missouri (Pflieger, 1975) and April to early July habitat) throughout its range. in Arkansas (Hill, 1968). My data based on Ar- kansas stocks are in accord with a late repro- Greensidedarter.-The reproductive season for ductive season (Fig. 6). The earliest ripe females Etheostomablennioides has been reported as late were 9 May; seven species from the same lo- March and early April in Missouri (Pflieger, calities were reproductive in April. I have also 1975) late April in Kentucky (Kuehne and Bar- obtained ripe females in late June supporting bour, 1983) and April and May in Michigan Hill's report of early July reproduction. (Winn, 1958). Pflieger and Winn suggested that greenside darters cease breeding earlier than Niangua darter.-Pflieger (1975) reported other darters in Missouri, Michigan, Kentucky spawning activities by Etheostoma nianguae in and Tennessee. My data support early repro- early April. My documentation of a ripe female duction of greenside darters with the Arkansas on 20 April (Fig. 6) is in general agreement with season in late Feb. to late March and the Mis- Pflieger and suggests extended spawning. souri season in April (Fig. 6). Numerous samples of darters in both regions had reproductive fe- .-The reproductive season for males of other species but non-reproductive fe- Etheostomacaeruleum has been reported as late male E. blennioides throughout May. March to May in Missouri (Pflieger, 1975), early April to early June in Michigan (Winn, 1958) Fantail darter.-The reproductive season for and April to June in Wisconsin (Becker, 1983). Etheostomaflabellare has been reported to be My Missouri data support the season reported April to June and perhaps July (Page, 1983) at by Pflieger (Fig. 6). My Arkansas data show various localities. It has been recorded as April spawning from late Feb. to late May. The re- and May in Missouri (Pflieger, 1975). My data productive season is two to three months in most support his conclusion with reproductive fe- areas. males recorded on many days during those two months (Fig. 6). DISCUSSION

Speckled darter.-The reproductive season for Darters in most geographic regions treated Etheostomastigmaeum has been reported to be here have breeding seasons extending over sev- April in Kentucky and Tennessee (Winn, 1958), eral months. The cumulative data suggest that April and May in Kansas (Cross, 1967) and Mis- in stenothermal environments in Texas three souri (Pflieger, 1975) and late March to early darter species (E. lepidum, E. grahami and E. April in many localities (Howell, 1978). My data fonticola) are reproductive all year. In addition, for Arkansas stocks suggest that the inclusive fertilized eggs have been obtained over inter- range listed for the several locations is correct vals of 273 (E. lepidum), 203 (E. spectabile), 150 (Fig. 6). I have records of ripe females from 20 (P. sciera), 143 (P. caprodes), 100 (E. proeliare), March to 16 May and numerous additional rec- 99 (P. shumardi), 96 (E. caeruleum), 88 (E. gra- ords of clear eggs from females on other dates hami), 86 (E. punctulatum), 76 (E. chlorosomum) including some as late as 30 May. and 71 (E. gracile, E. radiosum, E. tetrazonum,E. stigmaeum and E. flabellare) days. Except for E. Arkansas saddled darter.-The only data avail- blennioides and E. juliae, the other species have able on the reproductive season for Etheostoma data for fewer than five days of reproduction. euzonumare that provided by Hubbs and Strawn If eggs have been removed from a female on (1957b) on fertilized eggs obtained in May. I only one occasion, obviously, only one repro- now have data on fertilized eggs from 30 March ductive day can be reported. Documentation of to 12 May (Fig. 6). prolonged spawning must follow study over a protracted interval that is not available for the Missouri saddled darter.-The reproductive sea- species with fewer than ten known days of re- son for Etheostomatetrazonum has been reported production. Thirteen species are known to be HUBBS-DARTER REPRODUCTIVE SEASONS 65

reproductive for more than two months. Each The breeding season for logperch in Texas species has been shown to have most females in is substantially shorter than that for either E. reproductive cycle throughout that interval; lepidum or E. spectabile. The cumulative season thus, individual females are reproductive for is 143 days and individual populations are known that time interval. Data on E. lepidum and E. to spawn for up to 140 days. The major differ- spectabilesuggest that interbrood intervals range ence is in the date of first spawning (Dec. or from 4 to 10 days dependent upon stream tem- Jan. vs Oct., Nov. or Dec.) and most Texas perature. Most streams are at 15-20 C during populations start spawning at the time of short the reproductive season-at those tempera- daylength or shortly after. There is little dif- tures interbrood intervals are 5-10 days. ference in the date of reproductive termination Therefore, females of the 13 species have the in eurythermal populations. The breeding sea- potential of a minimum of six spawnings an- son for logperch in Arkansas (62 days) is shorter nually; if the stream temperatures are warmer, than that for Texas populations and again the the minimum number of spawnings would be majority of that difference is in reproductive 12. The maximum number of clutches per year initiation (Feb. vs March) not termination (mid- is 73. May vs early June). The spawning season in Mis- The spawning season for central Texas dart- souri seems displaced seasonally but Pflieger's ers is long. The recorded season for E. lepidum (1975) report of April breeding suggests that is 273 days and ripe eggs have been obtained the actual season is April through early June, periodically during the other 93 days. The doc- again different dates of spawning initiation and umented length of the season in individual comparable termination dates. Winn's (1958) stenothermal environments is 246 (S. Guada- report of April to early June reproduction in lupe), 215 (Mtn. Home) and 211 (S. Concho) southern Michigan is in accord with a delay in days and that of extensively sampled euryther- initiation of spawning but little difference in mal environments 164 (Junction), 167 (San Saba) termination. and 162 (Ingram) days. Thermal stability plays Geographic comparisons of spawning season a substantial role in the length of the repro- within a species are in accord with a pattern of ductive season. Whenever the temperature be- longer seasons in the south than in the north. comes warm (=ca 24 C) spawning activity slows Additionally, comparisons among geographi- or ceases. Each eurythermal population shows cally restricted species show the same pattern, spawning season initiation during declining with Texas species often spawning for most of daylength. the year, and those from Arkansas or Missouri The breeding season for E. spectabilein Texas spawning for two to three months. Few reports is quite similar to that of eurythermal popula- for more northern species are for more than tions of E. lepidum. The summed reproductive two months of spawning. season is 203 days and individual populations The reproductive seasons for darters vary ex- are known to be reproductive for at least 147 tensively from essentially continuous spawning (Ingram) to 187 (N. Guadalupe) days. The ini- in stenothermal waters in southern locations tiation of reproduction is consistently during through prolonged winter/spring spawning in periods of decreasing daylength. Marsh's (1980) eurythermal waters in southern locations to rel- documentation of extended spawning in artifi- atively brief spawning at northern latitudes. cally cooled waters shows that temperature has Much effort has been focused on the causal fac- an effect on length of spawning season. No sub- tors for initiation of spawning but much less stantial shift in spawning season with latitude is attention has been directed to causal factors for apparent among Texas populations. There is a termination of spawning. Especially for organ- suggestion of a shift between the northernmost isms with substantial geographic variation in Texas sample (Lampassas) and the others as it spawning seasons it is possible that different starts last and ends last. The Arkansas popula- mechanisms could apply. The variations in the tions have markedly shorter seasons (ca 100 days) reproductive seasons for E. spectabile may pro- than the Texas populations. The majority of vide insight into reproductive cues. Although the change is at spawning initiation (Feb. vs Nov.) annual clocks may play a role in reproductive with little change in last spawning (early June stimulation, Marsh's (1980) demonstration of a vs mid-May). The more northern populations change of the date of termination associated continue the trend with shorter spawning sea- with an artificial change in environmental tem- sons that start in March (Pflieger, 1975; Winn, perature suggests, at least, that termination is 1958) and end in early June. not totally dependent on an endogenous rhythm. 66 COPEIA, 1985, NO. 1

16

25- 15 :N

14 0 ~ 15 13-

Edwards Plateou Lompossos 12

5 - N.W. Arkansas I / I Kerrvilne,TX aompa sos TX ' 0 ' W.Central Missouri T I0-'/ s --eFayettevitle,ARK - Urbono,ILL II0 : -Jefferson City, MO Adrian,MI :/ \:,';->.:';;-sUrban , ILL -5 Adrion,MT HigginsA Loke,MI 9 v.j HHiggins Lake, MI I _ I . I t t - I I I I { ti I I : I J 'F'M' A'M J'A S O N D J J:FM AM J A SO N D J 8. Annual for seven localities de- Fig. 7. Annual thermal cycles for seven locations Fig. light cycles from The localities are derived from NOAA (1974). The localitiesare those rived Smithsonian(1896). in the most studied for darter closest to the regions most closely studied for darter regions closely reproduc- tion in the central United States. Known dates for reproductionin the centralUnited States.Known dates Etheostoma initiation and ter- for Etheostomaspectabile reproductive initiation and spectabilereproductive mination are arrows. terminationare designated by arrows. designated by

The commonly discussed environmental cues Those data reflect the last known date for are light and temperature. In addition, food spawning and if a stimulus period is considered, supplies must be sufficient to provide metabo- the most northern population has the lowest lites for gamete production; the wide scatter spawning thermal maximum and the slowest = (standard deviation 20 + eggs) in fecundity warming. Thus, a time lag between the cue to of equal-sized females recorded for E. lepidum stop spawning and the actual stop would have and E. spectabile(Hubbs et al., 1968) is best ex- the greatest thermal consistency among the cue plained by variations in food supplies. temperatures in northern populations. The ma- The air temperature cycles (Fig. 7) for the jor exception in termination of intraspecific various locations inhabited by E. spectabileshow spawning seasons is the later time for P. caprodes the expected colder weather in more polar lo- in northern Michigan; however, it is this area cations. In addition, the summer temperatures that has much reduced mid-summer warming. are more similar than are the winter tempera- The pattern of light cycles and spawning in- tures. The major exception is that summers in tervals shows the opposite pattern (Fig. 8). northern Michigan (Higgins Lake) are about 3 Spawning initiation is 9 hrs 10 min daylight to C colder than in southern Michigan (Adrian) 12 hours daylight. In this instance the short day- whereas the winters are only about 4 C colder. length is for southern locations and any stimulus The same comparison between Lampassas, would involve a prior interval with longer day- Texas, and NW Arkansas shows differences of light; the longer daylight figures are for north- 2 C and 5 C. The temperatures for first spawn- ern populations and for these the preceding in- ings vary widely (17, 9.5, 5, 6, 5.5 C) from south tervals are for less daylight. In effect, daylength to north. The distinction between southern and for initiation of reproductive stimuli is likely to northern populations must be even greater than be reasonably concordant among the popula- suggested by spawning dates as the stimulus for tions. The major difference would be that for spawning must precede actual egg deposition. southern populations it occurs in decreasing Thus, in the areas with the warmer tempera- daylength and it would be likely to involve in- tures an earlier date would have even warmer creasing daylength in northern populations. conditions and the areas with the colder tem- There is somewhat less variation between peratures would be even colder than these of daylength of last spawnings between 13 hrs 40 the actual spawning date. min daylight in the south and 15 hrs 10 min in The pattern of environmental temperatures the north. Although the variation of daylength for spawning termination shows much more at end of spawning is not dissimilar to those for similarity with all between 21.5 and 23.7 C. spawning initiation, they also correspond well HUBBS-DARTER REPRODUCTIVE SEASONS 67

with season of the year. More critically, the sub- BURR, B. M., AND R. L. MAYDEN.1979. Records of stantial change in date of spawning termination fishes in western Kentucky with additions to the known fauna. Trans. Acad. Sci. 40:58-67. with thermal change (Marsh, 1980) suggests a Ky. role of in the , AND L. M. PAGE. 1978. The life history of major temperature determining the end of the season. An that cypress darter, Etheostoma proeliare, in Max spawning hypothesis Illinois. Ill. Nat. Hist. Surv. Biol. Notes 106. a role in ter- Creek, temperature plays major spawning CLOUTMAN, D. G. 1980. Etheostoma mination is the punctulatum supported by year-round spawn- (Agassiz). Stippled darter, p. 684. In: Atlas of North ing of populations (E. lepidum, E. grahami, E. American freshwater fishes. D. S. Lee et al. (eds.). fonticola) inhabiting stenothermal waters. Sim- North Carolina State Mus. Nat. Hist. ply, some factor has turned on reproduction COLLETTE,B. B. 1962. The swamp darters of the and a thermal turn-off would not occur as fe- genus Hololepis(Pisces, ). Tulane Stud. Zool. males would not experience that level of ther- 9:115-211. F. B. 1967. Handbook of fishes of Kansas. mal extreme during their entire lives. CROSS, The on E. Hubbs and Misc. Publ. Mus. Nat. Hist. Univ. Kansas 45. experiments lepidum by R. F. 1969. A of the Strawn should be reevaluated in this DENONCOURT, systematic study (1957a) darter, Percina evides and context. females continued to de- gilt (ordan Copeland) Reproductive (Pisces, Percidae). Unpubl. PhD Diss., Cornell Univ. posit eggs throughout their lives at varied day- FORBES,S. A., AND R. E. RICHARDSON.1908. The lengths between 0 and 24 hours light. No dif- fishes of Illinois. Illinois Nat. Hist. Survey, Urbana. ference in egg production could be correlated GALE,W. R., ANDW. G. DEUTSCH.1985. Fecundity with daylength. However, when the water tem- and spawning frequency of captive tessellated dart- perature was raised from 20 to 23 C, there was er-fractional spawners. Trans. Amer. Fish. Soc. 114. a marked drop in reproductive activity. This HARRELL,H. L. 1980. Etheostoma correlates well with the observed temperatures grahami (Girard) Rio Grande darter, 652. In: Atlas of North Amer- in the environment when reproduction ceases p. ican freshwater fishes. D. S. Lee et al. (eds.). North (in Texas eurythermal environments they are Carolina State Mus. Nat. Hist. the same as for E. the spectabile). Additionally, HILL, L. G. 1968. Inter- and intrapopulational vari- above which E. temperature (24 C) lepidum egg ation of vertebral numbers of the yoke darter, Ethe- incubation success is low (Hubbs, 1961; Hubbs ostomajuliae. Southwest. Nat. 13:175-191. et al., 1969), is quite similar to that at which HOWELL,W. M. 1978. and distribution reproduction ceases. of the period fish, Etheostomastigmaeum, with the validation and redescription of Etheostomadavidsoni Hay. Unpubl. PhD Diss., Cornell Univ. ACKNOWLEDGMENTS HUBBS,C. 1961. Developmental temperature toler- ances of four etheostomatine fishes occurring in The field studies for this research were done Texas. Copeia 1961:195-198. under a series of NSF grants including GB 3206. . 1967. Geographic variations in survival of I also thank officials of the states of Arkansas, hybrids between etheostomatine fishes. Bull. Texas Mem. Mus. 13. Missouri, Oklahoma and Texas for numerous 1971. Competition and isolation mechanisms collecting permits. I have benefited from dis- in the Gambusiaaffinis x G. heterochirhybrid swarm. cussions with Lawrence M. Page, Edie Marsh Ibid. 19. and Gary P. Garrett on darter reproduction. 1982. A checklistof Texas freshwaterfishes. This manuscript has been improved by reviews Texas Parks and Wildlife Tech. Series 11. of a preliminary draft by Dr. Page and Robert 1983. Handbook of darters (review). Copeia A. Kuehne. A large number of associates have 1983:581. contributed to the field and laboratory work, ,AND C. BRYAN. 1975. Ontogenetic rates and tolerances of the channel most notably Alex Peden, Neal Armstrong, darter, Hadropterus cope- landi. Texas Sci. 24:623-625. William F. Hettler, F. Douglas Martin and Kirk J. Strawn. , ANDP. S. MARTIN. 1965. Effects of darkness on egg deposition by Etheostoma lepidum females. Southwest. Nat. 4:302-306. LITERATURE CITED ,ALEX E. PEDEN AND MICHAEL M. STEVENSON. 1969. The developmental rate of the greenthroat BECKER,G. C. 1983. Fishes of Wisconsin. Univ. Wis- darter, Etheostoma lepidum. Amer. Midi. Nat. 81: consin Press. 182-188. BRAASCH,M. E., AND P. W. SMITH. 1967. The life , AND J. PIGG. 1972. Habitat preferences of historyof the sloughdarter, Etheostoma gracile (Pisces, the harlequin darter, Etheostoma histrio, in Texas Percidae). Ill. Nat. Hist. Surv. Biol. Notes 58. and Oklahoma. Copeia 1972:193-194. 68 COPEIA, 1985, NO. 1

, M. M. STEVENSONAND A. E. PEDEN. 1968. SCALET, C. G. 1973. Reproduction of the orange- Fecundity and egg size in two central Texas darter belly darter, Etheostomaradiosum cyanorum (Osteich- populations. Southwest. Nat. 13:301-323. thyes, Percidae). Amer. Midi. Nat. 89:159-165. , AND K. STRAWN. 1957a. The effects of light SCOTT, W. B., AND E.J. CROSSMAN.1973. Freshwater and temperature on the fecundity of the green- fishes of Canada. Bull. Fish. Res. Bd. Canada 184. throat darter, Etheostomalepidum. Ecology 38:596- SCHENCK, J. R., AND B. G. WHITESIDES. 1977. Re- 602. production, fecundity, sexual dimorphism and sex ,AND . 1957b. Survival of Fl hybrids ratio of Etheostomafonticola (Osteichthyes, Perci- between fishes of the sub-family . dae). Amer. Midi. Nat. 98:365-375. J. Expt. Zool. 134:31-60. SMITH,P. W. 1979. The fishes of Illinois. Univ. Ill. KUEHNE,R. A., AND R. W. BARBOUR. 1983. The Press, Urbana. American darters. Univ. Press of Kentucky. SMITHSONIANINSTITUTION. 1896. Meteorological LACHNER,E. A., E. E. WESTLAKEAND P. S. HANDWERK. tables. Smith. Misc. Coll. 1950. Studies on the biology of some percid fishes STARNES, W. C. 1977. The ecology and life history from western Pennsylvania. Amer. Midi. Nat. 43: of the endangered snail darter, Percina (Imostoma) 92-111. tanasi Etnier. Tenn. Wildl. Res. Agency Tech. Rep. LAGLER, K. F., J. F. BARDACH, R. R. MILLERAND D. 77-52. R. M. PASSINO. 1977. Ichthyology.John Wiley and STRAWN, K. 1956. A method of breeding and raising Sons. three Texas darters. Part II. Aquarium J. 27:11, MARSH,E. 1980. The effects of temperature and 13, 14, 17, 31, 32. photoperiod on the termination of spawning in the . 1961. A comparison of meristic means and orangethroat darter (Etheostomaspectabile) in central variances of wild and laboratory-raised samples of Texas. Tex.J. Sci. 32:129-142. fishes, Etheostomagrahami and E. lepidum (Percidae). MATHUR, D. 1973. Food habits and feeding chro- TexasJ. Sci. 13:127-159. nology of the black-banded darter, Percina nigro- , AND C. HUBBS. 1956. Observations on strip- fasciata (Agassiz), in Halawalee Creek, Alabama. ping small fishes for experimental purposes. Copeia Trans. Amer. Fish. Soc. 102:48-55. 1956:114-116. MILLER, R.J., AND H. W. ROBISON. 1973. The fishes STEVENSON, M. M. 1971. Percina macrolepida(Pisces, of Oklahoma. Oklahoma State University Press. Percidae, Etheostomatinae), new percid fish of the MORRIS, M. A., AND L. M. PAGE. 1981. Variation in subgenus Percina from Texas. Southwest. Nat. 16: western (Pisces: Percidae), with descrip- 65-83. tion of a new subspecies from the Ozarks. Copeia THOMAS, D. L. 1970. An ecological study of four 1981:95-108. darters of the genus Percina (Percidae) in the Kas- MOYLE, P. B. 1976. Inland fishes of . Univ. kaskia River, Illinois. Ill. Nat. Hist. Surv. Biol. Notes Calif. Press. 70. NIKOLSKI, G. V. 1963. The ecology of fishes. Aca- THOMPSON, B. A. 1978. Logperches of southeastern demic Press. United States (Etheostomatini, Percina). ASB Bull. NOAA. 1974. Climates of the states. Vols. 1 and 2. 25:27. NOAA. WILLIAMS,J.D. 1975. Systematics of the percid fishes PAGE, L. M. 1983. Handbook of darters. TFH Pub- of the subgenus Ammocrypta,genus Ammocrypta,with lications. descriptions of two new species. Bull. Ala. Mus. Nat. , AND P. W. SMITH. 1970. The life history of Hist. 1. the dusky darter, Percina sciera, in the Embarras WINN, H. E. 1958. Comparative reproductive be- River, Illinois. Ill. Nat. Hist. Surv. Biol. Notes 69. havior and ecology of fourteen species of darters , AND . 1971. The life history of the (Pisces-Percidae). Ecol. Monogr. 28:155-191. slenderhead darter, Percina phoxocephala,in the Em- barras River, Illinois. Ill. Nat. Hist. Surv. Biol. Notes 74. DEPARTMENT OF ZOOLOGY, THE UNIVERSITY OF PFLIEGER, W. L. 1975. The fishes of Missouri. Mis- TEXAS AT AUSTIN, AUSTIN, TEXAS 78712. souri Department of Conservation. Accepted 27 April 1984.