From the Yucatán Peninsula, Belize and Guatemala Revista De Biología Tropical, Vol

Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica

Brandorff, Gerd-Oltmann Distribution of some Calanoida (Crustacea: Copepoda) from the Yucatán Peninsula, Belize and Guatemala Revista de Biología Tropical, vol. 60, núm. 1, marzo, 2012, pp. 187-202 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Distribution of some Calanoida (Crustacea: Copepoda) from the Yucatán Peninsula, Belize and Guatemala

Gerd-Oltmann Brandorff Georg-Gröning-Str. 29 A, D-28209 Bremen, Germany; [email protected]

Received 07-iii-2011. Corrected 22-vii-2011. Accepted 29-viii-2011.

Abstract: Southern Mexico and Central America have many water bodies of different morphology and water chemistry with an interesting zooplankton fauna, originating from North or South America. A set of 63 samples, taken in 2005 and 2008, from water bodies of the Yucatan Peninsula karst, Belize and Guatemala, were studied for the content of calanoid copepods. Old and recent literature was used to determine animals to species level. Drawings were prepared with a microscope and a camera lucida. A total of 32 samples with totally six species contained calanoid copepods: one estuarine pseudodiaptomid and five freshwater diaptomids. Pseudodiaptomus marshi was found at different salinities. It is confirmed that the commonest diaptomids in the Yucatan Peninsula are Arctodiaptomus dorsalis and Mastigodiaptomus nesus. The former was also recorded from Lake Amatitlan. Mastigodiaptomus nesus is as widespread as A. dorsalis but it is absent from the Lake Peten area in Guatemala. Mastigodiaptomus reidae was found in two shallow habitats, these specimens differ from those from the type locality by having a set of peculiar large spine-like processes on the last thoracic and the urosome segments of the females. Leptodiaptomus siciloides was found only in Lake Ayarza with high salinity. Prionodiaptomus colombiensis occurred in the highlands of Guatemala in Lago de Güija and in the Peten area in Laguna Sacpuy. We contributed with our occurrence records to a better knowledge of the geographic distribution of some calanoid copepods. Morphological findings in some species are of value for taxonomic differentiation between species. Rev. Biol. Trop. 60 (1): 187-202. Epub 2012 March 01.

Key words: Central America, zooplankton, calanoid copepods, Pseudodiaptomidae, Diaptomidae, geographic distribution.

Mexico, Central America and the Carib- genera, and one (Diaptomus s.l.) with unclear bean, as a bridge between North and South taxonomical classification. From the tropical America, host many species from two distinct Yucatan Peninsula and Guatemala three genera biogeographical regions, the Nearctic and Neo- with seven species and one pseudodiaptomid tropical. It is also called Neotropical-Caribbean copepod are known. (NCAR) (Dussart & Defaye 2001). Zooplank- Decades ago Dussart et al. (1984) consid- ton represents an important community to ered the Central American region as “well stud- transfer matter and energy between producers ied” for the freshwater copepods. Although, and consumers in food webs, thus shaping new species have been described, especial- the pelagic ecosystem. Copepods make up a ly from Mexico (Suárez-Morales & Elías- major portion of the biomass and productivity Gutiérrez 2000), Elías-Gutiérrez et al. (2008a) of freshwater ecosystems (Williamson & Reid showed that there are still many overlooked/ 2001). Nine genera with 26 species of diaptom- cryptic species of Diaptomidae in the region id copepods (Crustacea) are registered (Suárez- to be morphologically described. But, there Morales et al. 2005), five genera from the are still many water bodies to be sampled, for Nearctic, one from South America, two NCAR instance, Yucatan cenotes and lakes in adjacent

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 187 areas (Alcocer & Bernal-Brooks 2010). So it Environmental Geology of the University of was a good opportunity to look at the samples Braunschweig, Germany sampled freshwater from the Yucatan Peninsula, Belize and Gua- habitats in the Yucatan lowlands of Mexico, temala that were presented to me and to see Belize and Guatemala (Pérez et al. 2010). The what calanoid species they contain. In this aim of their project was to use ostracods from contribution the results of the extensive sam- the Yucatán Peninsula as indicators of past pling of calanoid copepods of inland waters are environmental and climate change. Sampled presented, and comparative morphological and sites included: lakes, sinkholes, brackish water distributional comments about the species are lagoons, rivers in the lowlands, and lakes in the provided herein. Guatemalan highlands (Fig. 1, Tables 1, 2). Water samples to determine physico-chem- MATERIALS AND METHODS ical variables as temperature, pH, conductivity and oxygen were taken with a Ruttner-sampler In 2005 and 2008 Burkhard Scharf, Liseth and measured with a WTW multisensor equip- Pérez and co-workers from the Institute of ment. Qualitative samples of plankton in the

Fig. 1. Sampled water bodies in 2005 and 2008, their numbers and geographical position.

188 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012

X Ls

X X Pc

X X X X X X X X Ad

X Pm 630 215 465 568 206 232 850 232 283 238 285 187 179 292 568 1 772 4 310 1 019 [µS/cm] Conductivity ] 2 7 2 11 45 14 2.5 1.1 1.6 2.9 3.5 3.6 3.8 5.6 645 126 100 15.2 0.02 [km Surface area 4 2 6 8 3 23 15 21 80 22 10 38 65 35 40 [m] 340 165 250 Depth 4 TABLE 1 TABLE 75 115 114 115 117 433 165 219 587 148 122 132 134 146 1200 1560 1414 [m asl] Altitude W 90°32’58.6’’ 89°06’39.0’’ 91°09’16.6’’ 89°51’14.3’’ 89°32’50.2’’ 90°12’29.2’’ 89°37’53.7’’ 89°24’29.7’’ 89°41’39.2’’ 90°08’10.2’’ 89°44’27.4’’ 89°40’30.9’’ 90°00’52.2’’ 90°22’10.3’’ 90°25’21.6’’ 90°10’03.6’’ 90°10’48.9’’ 90°09’36.2’’ Coordinates N 14°26’03.7’’ 15°27’18.0’’ 14°43’57.9’’ 17°00’21.3’’ 14°14’53.0’’ 17°04’09.5’’ 16°58’16.3’’ 17°03’54.9’’ 14°13’01.1’’ 14°26’02.0’’ 16°39’18.2’’ 16°58’38.2’’ 16°58’32.4’’ 16°56’51.0’’ 16°55’08.3’’ 16°20’15.1’’ 16°25’10.3’’ 16°31’31.4’’ Name of waterbody List of sampled lakes, their geographic position and some morphometric, limnochemical data and found freshwater calanoid copepod species data and found freshwater calanoid limnochemical position and some morphometric, geographic lakes, their List of sampled Amatitlán I zabal Atitlán Petén I tza Güija Perdida Macanché Yaxhá Atescampa Ayarza Oquevix Salpeten Sacpuy La Gloria San Diego Las Pozas Petexbatun El Rosario 1 2 3 4 5 19 20 21 22 23 25 26 28 29 30 32 33 34

I D-Nr.

highlands

Guatemalan lowlands Guatemalan Guatemala Guatemala Area

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 189

X X X X X X X X X X Mn

X X X X X Ad

X X Pm Ls Leptodiaptomus siciloides 330 774 754 488 139 146 174 241 474 127 1221 1231 1 411 2 720 1 715 1 481 2 060 2 350 1 213 [µS/cm] Conductivity ] 2 2 5 2 51 23 3.1 1.5 3.9 8.5 5.1 0.9 2.9 1.2 0.1 0.25 0.25 0.02 0.35 [km Surface area ) 4 2 3 8 3 3 – 3 3 5 6 8 – 11 16 10 14 20 0.6 [m] Depth Pc Prionodiaptomus colombiensis , 1 1 1 2 1 1 1 2 3 2 7 38 91 90 74 52 57 69 107 [m asl] Altitude TABLE 1 ( Continued TABLE W 88°25’37.5’’ 88°18’42.4’’ 88°23’22.4’’ 88°31’49.1’’ 88°26’05.6’’ 88°10’29.3’’ 88°03’15.2’’ 88°46’00.1’’ 87°38’10.8’’ 89°13’10.8’’ 89°00’41.8’’ 88°34’20.6’’ 89°05’13.9’’ 89°10’17.0’’ 90°06’45.4’’ 90°39’22.8’’ 90°28’47.3’’ 90°58’33.0’’ 87°44’19.2’’ Mn Mastigodiaptomus nesus , Coordinates N 18°22’11.8’’ 19°11’21.8’’ 18°30’49.8’’ 17°27’51.0’’ 18°39’53.3’’ 17°46’54.0’’ 18°02’44.3’’ 19°08’47.3’’ 19°28’27.2’’ 19°52’48.7’’ 20°38’52.5’’ 20°39’25.6’’ 18°24’03.2’’ 18°28’42.1’’ 18°41’40.7’’ 17°53’55.9’’ 18°02’48.9’’ 18°02’34.3’’ 20°29’40.2’’ Arctodiaptomus dorsalis , Arctodiaptomus Ad Name of waterbody List of sampled lakes, their geographic position and some morphometric, limnochemical data and found freshwater calanoid copepod species data and found freshwater calanoid limnochemical position and some morphometric, geographic lakes, their List of sampled Milagros Almond Hill Lagoon Bacalar Lagoon Crooked Tree Honey Camp Lagoon Nohbec Ocom Chichancanab Punta Laguna Yalahau San José Aguilar Sabanita Chacan-Bata Jobal San Francisco Mateos Chacan-Lara La Miseriosa Cayucon Coba 8 9 12 13 10 14 15 16 17 18 36 37 38 41 42 39 43 45 61

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lowlands

Mexican lowlands Mexican Belizean Belizean Area Pm Pseudodiaptomus marshi ,

190 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012

X X Mr

X X X Mn

X Ad

X Pm 793 958 643 949 720 183 168 244 192 192 4340 2 110 2 520 2 700 1 564 2 040 1 465 2 520 1 025 2 040 5 960 1 751 1 452 3 670 38 200 55 300 [µS/cm] Conductivity ] 2 ------28 7.2 2.3 0.03 0.03 0.06 0.01 [km <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 Surface area 1 1 2 1 1 1 1 4 7 47 25 45 --- 45 --- 1.5 1.5 1.5 0.5 2.5 3.2 1.5 1.5 1.5 0.5 11.1 Depth [m] Mastigodiaptomus cf. reidae 5 3 9 5 7 7 6 3 4 4 Mr 27 18 44 80 14 20 13 59 33 25 77 14 115 110 141 179 [m asl] Altitude TABLE 2 TABLE W 89°53’27,8” 90°11’00.3’’ 89°29’45.1’’ 88°24’56.0’’ 90°43’47.6’’ 88°15’52.5’’ 91°02’59.6’’ 90°28’39.1’’ 87°20’23.8’’ 88°08’30.8’’ 88°43’52.0’’ 89°52’34.5’’ 89°21’23.7’’ 90°17’39.8’’ 90°08’14.9’’ 88°29’18.2’’ 88°24’55.3’’ 88°22’57.4’’ 89°50’03.2’’ 89°35’53.3’’ 90°19’13.8’’ 90°07’04.5’’ 88°57’49.3’’ 88°45’30.8’’ 90°22’39.2’’ 89°18’01.9’’ Coordinates Mn Mastigodiaptomus nesus , N 16°58’27.3” 18°11’02.4’’ 16°38’11.6’’ 21°20’11.3’’ 19°27’58.3’’ 20°37’15.4’’ 19°12’41.6’’ 20°36’08.1’’ 17°56’55.4’’ 20°48’09.6’’ 21°25’04.3’’ 20°42’24.6’’ 20°41’23.0’’ 20°35’38.8’’ 18°38’33.0’’ 20°52’03.6’’ 17°18’17.8’’ 18°13’04.4’’ 18°13’36.5’’ 20°45’00.9’’ 21°05’27.6’’ 20°50’59.6’’ 20°51’22.2’’ 15°40’25.3’’ 17°18’59.8’’ 20°51’20.8’’ I xlú Cuba Subin Timul Loché Juárez Tekom Xlacah Rosada Chenhá Silvituc Belize 2 Belize Belize 1 Belize Jamolún Celestun Progreso Guerrero Ya’ax’ex Rio Dulce Candelaria Yokdzonot Little Belize Little near Oquevix Arctodiaptomus dorsalis , Ad Arctodiaptomus Peten de Monos Name of waterbody San Francisco Kana San I gnacio Chochola 7 6 11 63 48 47 46 60 62 44 58 57 31 40 25 59 56 27 35 55 50 52 54 24 51 49

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water bodies water Cenotes Rivers Ponds List of sampled waterbodies others than lakes, their geographic position and some morphometric, limnochemical data and found freshwater calanoid copepod species data and found freshwater calanoid limnochemical position and some morphometric, geographic waterbodies others than lakes, their List of sampled Coastal Aquatic Wetland environment Pm Pseudodiaptomus marshi ,

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 191 open water and the littoral were taken using RESULTS standard plankton net with 100µm mesh size, fixed and preserved in 4% formalin. In the lab- Tables 1 and 2, list species of calanoid copepods from the lakes and other water bod- oratory, adult calanoid copepods were sorted ies, like cenotes, ponds, coastal water bodies under an Olympus dissecting microscope, and and rivers, in Yucatan Peninsula, Guatemala identified to the species level with an Olympus and Belize, arranged according to Pérez et al CH 2 microscope. The following literature (2010). Only 32 of the 63 sampled water bod- was used: Light (1939), Bowman (1986), ies contained calanoid copepods. Fig. 1 shows Gutiérrez-Aguirre & Suárez-Morales (2000), the sampled water bodies, their numbers and Suárez-Morales & Elías-Gutiérrez (2000, geographical position and figure 2 shows the 2001), and Elías-Gutiérrez et al. (2008b). geographical distribution of the species found Drawings were prepared with the Olympus CH in the sampled area. Pseudodiaptomus marshi 2 microscope, and a camera lucida attached Wright, 1936 occurred in four water bodies to the microscope at 40x magnification. The with fresh to oligosaline waters at nearly sea salinity classification of the lakes follows level (Fig. 2c). Our specimens from Lago Iza- Cowardin et al. (1979). bal, Progreso Lagoon and Almond Hill Lagoon

Gulf of Mexico Gulf of Mexico

Mexico Mexico

Caribbean Caribbean Guatemala Belize Guatemala Belize Sea Sea

Paci c Honduras Paci c Honduras Ocean El Salvador A Ocean El Salvador B

Gulf of Mexico Gulf of Mexico

Mexico Mexico

Caribbean Caribbean Guatemala Belize Guatemala Belize Sea Sea

Paci c Honduras Paci c Honduras Ocean El Salvador C Ocean El Salvador D

Fig. 2. Distributional patterns of the calanoid copepods found in our samples. a) Arctodiaptomus dorsalis, records from other authors (X) (Brinson & Nordlie 1975, Deevey 1980, Suárez 1991, Flores-Nava 1994, Gutiérrez-Aguirre & Suárez-Morales 2001, Elías-Gutiérrez et al. 2008a, b). b) Mastigodiaptomus nesus, records from other authors (X) (Elías-Gutiérrez et al. 2008a). c) Pseudodiaptomus marshi (rectangles) and Prionodiaptomus colombiensis (triangles), records from other authors (X for P. marshi and V for P. colombiensis) (Brinson & Nordlie 1975, Gutiérrez-Aguirre & Suárez-Morales 2001, Elías- Gutiérrez et al. 2008a, b). d) Leptodiaptomus siciloides (triangles) and Mastigodiaptomus cf. reidae (rectangles), data from other authors (X for M. reidae) (Suárez-Morales & Elías-Gutiérrez 2000, Elías-Gutiérrez 2008b).

192 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 possess an anterolateral knob with many spines In a pool and a wetland area (Fig. 2d) and hairs on the right side of female’s genital animals were found that resemble very much somite (Fig. 3). Mastigodiaptomus reidae Morales & Elías- Arctodiaptomus dorsalis (Marsh, 1907) Gutiérrez, 2000. The two water bodies contain was found in 14 water bodies that lie below an fresh water and lie at an altitude of about altitude of 200m.a.s.l., and in Lago Amatitlán 100m.a.s.l. Morphologically there are some (Guatemala) at 1 195m.a.s.l. (Fig. 2a). One of differences between our specimens and M. reidae, especially in the female (Fig. 4 a,b). these 14 water bodies (Lago Salpeten, Guate- On the not well developed border between mala) is oligohaline. pediger four and five on the posterior surface Mastigodiaptomus nesus (Bowman, 1986) on each side of the dorsal process we found was in 13 water bodies in the Yucatán Penin- two long, flexible spine-like processes, their sula at altitudes below 100m.a.s.l. (Fig. 2b). Six length is about 25 and 30µm. The dorsal lakes of this cluster are oligohaline. A. dorsalis process or hump we found only in some and M. nesus co-occurred in three lakes. specimens. Spine-like processes on left and right thoracic wing are on the same position as described in M. reidae, but are larger (about 25µm) in our specimens. About this same length was found for the two spines on the anterior third of the long urosome. The caudal rami have an inner and outer setose margin. Other morphological features are as

Fig. 4. Mastigodiaptomus cf. reidae: a) last pedigerous

somite and genital somite of female, b) left P5 of female, c) Fig. 3. Pseudodiaptomus marshi, female last pedigerous last pedigerous somite and urosome of male, d) P5 of male, somites and urosome, scale bar 0.1 mm, arrow indicates e) part of right P5 of male (another specimen) showing the anterolateral knob. ornamentation of exopodite 1, scale bar 0.1mm.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 193 in M. reidae. The male (Fig. 4 c-e) does not differ from the description of M. reidae. Most morphological differences between M. reidae and our specimens are small and probably lie within the species range. Only the spine-like processes on the female’s pediger 4/5 were remarkable and never seen in other specimens. Normally on this position one can find short fine hairs in many species. The morphologic variability in the M. cf. reidae females was conservative into and between surveyed popu- lations (Oquevix pond and Jamolún wetland). Prionodiaptomus colombiensis (Thie- Fig. 6. Box plots of Arctodiaptomus dorsalis and baud, 1912) was found in two freshwater Mastigodiaptomus nesus and the conductivity of the bodies in Guatemala at altitudes of 433m.a.s.l. sampled water bodies. and 206m.a.s.l. (Fig. 2c), respectively. In our specimens from Lake Güija we found, on the females genital segment, a fold on the ventro- Conductivity ranges of the water bodies posterior region that is crescent shaped (Fig. 5, where A. dorsalis and M. nesus were found, are double-arrow). shown in figure 6. Leptodiaptomus siciloides (Lilljeborg, 1889 in De Guerne & Richard) was found only in one highland lake (Fig. 2d) at an altitude of DISCUSSION 1 414m.a.s.l. with oligosaline water. An interpretation of some limnological and morphological characteristics, physical and chemical properties of the sampled water bodies waters has already been published (Pérez et al. 2010). From the 63 samples, taken in 2005 and 2008, only 32 contained calanoid copepods. The reasons why nearly 50% of the samples had no Calanoida can be numerous: wrong season, inadequate water chemistry, not enough food, predation, among others. Nine out of the 32 water bodies with Calanoida were sampled before by other authors. From the sampled region, one pseudodiaptomid and nine diaptomid species were known. We found one pseudodiaptomid and five diaptomid species, none of them was new, but we could add some new occurrences. The taxonomy of the Mexican species of Diaptomidae have made great progress in the last decades (Elías-Gutiérrez et al. 2008b), but still some species complexes with over- Fig. 5. Ventral (a) and lateral (b) view of genital segment looked species, as in A. dorsalis and Mastigo- of female Prionodiaptomus colombiensis with crescent diaptomus albuquerquensis, have to be cleared shaped fold (arrow), scale bar 0.1mm. (Elías-Gutiérrez et al. 2008a). Therefore it

194 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 is worth to note morphological differences According to Schmitter-Soto et al. (2002) in some species. A. dorsalis is the dominant copepod in the central epicontinental portion of the Yucatan Pseudodiaptomus marshi (Wright, 1936): Peninsula and is substituted in other portion is a widespread species in estuaries on the by Mastigodiaptomus ssp. Neighboring, there Atlantic coast of Central and South America, are some isolated occurrences in Campeche extending South to Vitória, Espirito Santo, Bra- and a cluster in Tamaulipas. In Guatemala zil (20°16’S) (Pereira & Loureiro Fernandes A. dorsalis is widely distributed in the Petén 2000, Santos Silva 2008). Brinson & Nordlie district (Deevey et al. 1980, this study) and in (1975) reported it as its Pacific sibling P. cule- Lago Izabal and Río Dulce (Brinson & Nordlie brensis from Lago Izabal. 1975, this study). Isolated occurrences were The estuarine species P. marshi can live in found in Lago Lachuá near the Mexican border a wide range of salinity, from fresh water (Lago (Elías-Gutiérrez et al. 2008a). The presence Izabal) to salinities up to 42ppt (Medeiros et al. in Lake Amatitlán (this study) was a great 2006) in Northeastern Brazil. In the Yucatan surprise. But only two males and two females Peninsula, the distribution was found mostly were found. Nearly 100 years before (1910) well inside the coastline (e.g. Laguna Bacalar), Mastigodiaptomus albuquerquensis (Herrick, that might have resulted from stranding and 1895) and M. amatitlanensis (Wilson, 1941) subsequent adaptation to freshwater during were reported (Juday 1915, Wilson 1941). Only a marine regression in the Bacalar formation more intensive sampling can show, if there is a (Suárez-Morales 2003). real change in species and if the massive eutro- Our results of the anterolateral knob on phication of the lake favored A. dorsalis and the females genital somite in P. marshi, is in excluded the Mastigodiaptomus species. accordance with the material from Costa Rica and Belize (Walter 1989), for which the genital Mastigodiaptomus nesus (Bowman, somite of specimens from Central America is 1986): It is distributed in the Yucatán Penin- asymmetrical, whereas from West Indies and sula (Suárez-Morales 2003) and the Northern South America it is symmetrical. Also Elías- Caribbean Islands (Bowman 1986, Reid & Gutiérrez et al. (2008b) showed an asymmetri- Hribar 2006). In the Yucatán Peninsula it cal genital somite from Mexican specimens, shows a scattered distribution in the lowlands among others from Laguna de Bacalar. of México and Belize. We found it in coastal Arctodiaptomus dorsalis (Marsh, 1907): lagoons, lakes, ponds and cenotes, but it was Its core range lies in tropical and subtropi- also found in wells, cave pool, potholes and cal lowlands bordering the Gulf of Mexico cenote-like pits (Bowman 1986), the extremes and Caribbean Sea, from the Southern United are Blue Holes where it was caught in depths States through Central America and Northern up to 50m (Fosshagen & Iliffe 2003, 2004). In South America, plus the Greater Antilles (Reid the deep Minicenote it is present throughout the 2007). In North America it extends its range by year and seems to make horizontal migrations human agency. to the walls of the cenote to escape predation In the studied area, it has a somewhat (Cervantes-Martínez et al. 2005). It was found disjunct distribution: in the Eastern coastal at very different salinities: from fresh water zone of Quintana Roo, in the North-western (Laguna Cayucón), different higher salinities part of the State of Yucatán round the city of in the Yucatán Peninsula to marine salinities Mérida, the Petén area in Guatemala, and the (Fosshagen & Iliffe 2004). So M. nesus is South-East of Tabasco. A similar picture for called a halotolerant form. The biogeography the distribution in the Yucatan Peninsula is of this species during geological development drawn by Suárez-Morales (2003). He also dis- of Central America is given by Suárez-Morales cusses the dispersal through geological times. & Reid (2003).

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 195 Mastigodiaptomus cf. reidae Suárez- M. reidae were found (Elías-Gutiérrez et al. Morales & Elías-Gutiérrez, 2000: M. cf reidae 2008a). Our specimens came from the same as is M. reidae (Suárez-Morales & Elías- type of habitat and also from the lowlands of Gutiérrez 2000) are relatively large diaptomids. Yucatán; whether it is one of the Kohunlich All were from the same area and from shal- species or a different one must be proven. low ponds. Their body size can be explained Reid et al. (2002) pointed out that the habi- with the lack of fish as predators, which can- tats of M. reidae and M. maya will be exposed not survive in these water bodies that are in to great human pressure because of increasing part ephemeral. A similar phenomenon was tourism development in the archaeological observed in the Pantanal, Brazil with Argyro- sites. These species from this area and the diaptomus nhumirim (Reid 1997, Brandorff specimens from the pond near Lake Oquevix et al. 2011). should be considered vulnerable. The dorsal process or hump occurs in many species of different genera, mostly is Prionodiaptomus colombiensis (Thie- very variable and therefore has no taxonom- baud, 1912): This species was found only in ic value (Wright 1927, Suárez-Morales & two samples from Laguna Sacpuy in Guatemala

TABLE 3 Female/male body size ratio of different Mastigodiaptomus species from published literature

Mastigodiaptomus mm Ratio Author species Female Male M. albuquerquensis 1.77 1.58 1.12 Marsh 1907 M. purpureus 2.56 2.24 1.14 Marsh 1907 M. nesus 1.48 1.34 1.10 Bowman 1986 M. montezumae 1.59 1.43 1.11 Santos Silva et al. 1996 M. amatitlanensis 1.45 1.33 1.09 Wilson 1941 M. maya 2.38 2.18 1.09 Suárez-Morales & Elías-Gutiérrez 2000 M. reidae 1.54 1.56 0.99 Suárez-Morales & Elías-Gutiérrez 2000

Elías-Gutiérrez 2000). Spine-like processes and Lago de Güija. On previous reports about on left and right thoracic wing are on the same the zooplankton of this lake, only immature position as described by Suárez-Morales & Diaptomidae were found (Marsh 1931). In June Elías-Gutiérrez (2000), but are larger (about 1980 Mastigodiaptomus albuquerquensis (Her- 25µm) in our specimens. About the same rick, 1895) was found in Lago de Güija (Ser- length are the two spines on the anterior third ruya & Pollingher 1983), while in our sample of the long urosome. The caudal rami have an from the same lake, P. colombiensis was the inner and outer setose margin. Striking is the only planktonic diaptomid. low female/male ratio of 0.99 in M. reidae One of the tables summarizes the pub- (Table 3), since normally the ratio is greater lished records on which P. colombiensis have than one and lies between 1.01 and 1.24 (Cic- been found and therefore its actual known dis- chino et al. 2001). tribution (Table 4). The Northern most point is M. reidae has been found in a small pond in Mexico and the Southernmost in Colombia. in Chicaná, Campeche (Suárez-Morales & According to Suárez-Morales et al. (2005) it is Elías-Gutiérrez 2000); and also in a small pond a South American genus and the only one “that at Kohunlich, Quintana Roo, where two other has radiated through Central America and reach morphologically similar but distinct species of Mexico”. The water bodies altitudes varied

196 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 ] 2 1.15 2.54 1.14 0.32 3.45 14.3 24.2 Area 8.157 423.1 [km 4 18 10 13 10 65 30 27 43 80? 50? 50? 123 427 688 709 105 Altitude [m.a.s.l.] 2 5 3 13 10 2.9 Mean 2 4 25 91 30 Depth [m] 2.11 2.79 0.67 2.22 Max West 92°31’ 91°32’ 91°31’ 92°47’ 92°17’ 92°48’ 89°05’ 90°00’ 89°32’ 89°51’ 89°40’ 88°03’ 86°46’ 85°21’ 79°40’ 79°58’ 79°52’ 79°52’ 88°36’ 94.972 90.649 90.145 Longitude 9°11’ 9°02’ 8°43’ 9°18’ North 11°35’ 17°52’ 17°47’ 17°45’ 18°23’ 18°23’ 18°17’ 16°58’ 15°16’ 14°14’ 13°57’ 13°59’ 13°18’ 15°46’ 15°43’ 16.714 15.951 15.776 Latitude Type shallow lake shallow lake shallow lake pond pond shallow lake pond shallow lake lake shallow lake deep shallow lake deep lake shallow lake lake pool TABLE 4 TABLE Environment Name icario Lagoon Laguna Matillas L. V Lechugal lagoon Lechugal km 51 pond 11 km 51 Laguna 3 Brazos Matias Romero Laguna Sacpuy Pond near Lachua 2 Chajmaic To Los Amates Lago de Güija Lake Ahuachapan (Laguna Llano Ahuachapan Lake del Espino?) Lake Chalchuapa Lake Olomega Lago Nicaragua at La Ceiba Reservoir Comacho Rio Trinidad Gatun Lake Brazos Brook Reservoir Pools near Puerto Barrios Author Gutiérrez-Aguirre & Suárez- Gutiérrez-Aguirre Morales 2000 Gutiérrez-Aguirre & Suárez- Gutiérrez-Aguirre Morales 2001 Elías-Gutiérrez et al. 2008a Elías-Gutiérrez Elías-Gutiérrez et al. 2008b Elías-Gutiérrez this study Elías-Gutiérrez et al. 2008b Elías-Gutiérrez Marsh 1931 Herbst 1960 Marsh 1919 no data Marsh 1913 Dodds 1926 Juday 1914 and additional data about the water bodies of Prionodiaptomus colombiensis and additional distribution about the geographical Data from the literature Mexico Panama Country Honduras Nicaragua Guatemala Costa Rica El Salvador 1 2 3 4 5 6 7 8 9 11 12 13 14 15 16 18 17 19 20 21 22 10 No

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 197 ] 2 50 Area [km 800ha 5 11 50 80 50 70 30 100 100 520 2070 2540 2630 Altitude [m.a.s.l.] 2 1.5 mud from dried Mean 8 3 12 Depth [m] 5.5 Max West 73°56’ 73°45’ 74°12’ 74°45’ 74°10’ 74°29’ 74°20’ 69°10’ 67°23’ 67°37’ 67°35’ 65°10’ 62°52’ Longitude 4°30’ 5°26’ 4°34’ 9°20’ 9°14’ 7°35’ 8°08’ 8°30’ 9°56’ North 11°00’ 10°11’ 10°31’ 10°35’ Latitude Type ) eutrophic shallow lake eutrophic pools marsh shallow lake Floodplain lake with lots lake Floodplain vegetation of aquatic pond permanent waters in marsh permanent Hypertroph shallow lake swamp pool pool ephemeral pond ephemeral Environment TABLE 4 ( Continued TABLE Name III , Peninsula de Paria, Laguna de Ubaque Laguna Fuquene marsh at Fundación marsh at Barranquilla Laguna de la Herrera Cienaga de Pigiño Cienaga Pond ‘Las Bolas’ Hacienda Pond ‘Las Bolas’ “Universidad de Antioquia” Esteros de Mantecal Zuata reservoir near Cagua Zuata Estero de Camaguan shady pool, km 245, road No 2 charca, near Unaré river at Clarines charca, Palmares Sucre Author Thiebaud 1912; Gaviria 1989 Thiebaud Gaviria 1994 Pearse 1915 Kiefer 1956 (Franke 1989) Gaviria pers. com. 2009 Montiel & Zoppi de Roa 1979 Montiel Dussart 1984 Torres & Zoppi de Roa Torres and additional data about the water bodies of Prionodiaptomus colombiensis and additional distribution about the geographical Data from the literature enezuela Country Colombia V 23 24 25 26 27 28 29 30 31 32 33 34 35 No

198 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 from nearly sea level (4m) to 2 540m in the highlands. The lake lies at an altitude of Colombian Andes, but more than 70% lie at an 1 414m.a.s.l., maximum depth is 250m and altitude between 0 and 100m. conductivity between 1 772 and 1 844µS/cm. P. colombiensis was found in water bod- A comparable habitat is Lago Ilopango in El ies of very different sizes: from small ponds Salvador, a lake of volcanic origin at an altitude and marshes to lakes of over 8 000km2. Most of 490m.a.s.l., with a maximum depth of 215m samples were from small, shallow lakes, ponds and conductivity between 1.45 and 2.09mS and marshes. Water bodies with a greater sur- (Gophen & Walline 2005). There L. siciloi- face area mostly were shallow and with much des was found by Juday (1915), present at all aquatic vegetation, like Lake Güija (Guatema- depths, but most abundant between 10 and la/El Salvador). In the large Lake Nicaragua P. 25m, together with numerous protozoan Tintin- colombiensis was taken from the shore (Herbst nidae. Also present was the euryhaline rotifer 1960, Hartmann 1959). Nearly all records indi- Hexarthra fennica; it was also reported from cate that P. colombiensis is a species living in Lago Chamico in El Salvador (Marsh 1931), a small pools or ponds, in vegetated lakes or on deep volcanic lake. These lakes in Guatemala the shore of lakes. This is confirmed by Dodds and El Salvador seem to be the southernmost (1926) from Gatun Lake (Panama), where it points of the L. siciloides geographical dis- was found in shores and ponds, but always tribution. The geographical range covers the in small numbers, except in a very dirty and USA and Southern Canada (Suárez-Morales et plant grown pond between Gamboa and Sum- al. 2005). In Wisconsin (Torke 2001) L. siciloi- mit, where it was the most abundant species. des is fairly common in permanent ponds, In Gatun Lake itself Diaptomus s.l. gatunensis lakes and reservoirs. It prefers more eutrophic Marsh, 1913 is the abundant diaptomid (Zaret habitats and this selection let it expand to new & Suffern 1976). Also Marsh (1931) stated that habitats due to cultural eutrophication of many for P. colombiensis “its preferred habitat is in lakes and ponds in this state. shallow water or pools”. In summary, P. colom- In Mexico it is widely distributed in the biensis dwells in shallow lakes, in the littoral states of Aguascalientes, Puebla, Coahuila and and/or within the macrophytes. Sonora (Suárez-Morales et al. 2005, Elías- In describing the species of Prionodi- Gutiérrez et al. 2008b). In Aguascalientes state aptomus incarum from Peru, Cicchino et al. L. siciloides was found at altitudes between 1 (2004) showed on the female genital segment 850 and 2 010m.a.s.l., in permanent and non- “ventroposterior region with very conspicuous permanent water bodies, with different surface semicircular fold with adjacent wrinkled area areas and maximum depth (Dodson & Silva- medially, reaching posterior margin of genital Briano 1996). Other limnological data were not double-somite”. They pointed out that also reported but probably cover a wide range. specimens from Venezuela of P. colombiensis In two of the sampled water bodies, A. have this morphological peculiarity. In our dorsalis and M. nesus co-occurred (lake San specimens it is crescent shaped. Further studies Francisco de Mateos, Mexico and Crooked have to show if it is characteristic for the genus Tree Sanctuary Lagoon, Belize); the same Prionodiaptomus and its function. A good co-occurrence was reported from Minicenote overall morphological description and varia- (Elías-Gutiérrez et al. 2008b). Species of the tion is given by Gutiérrez-Aguirre & Suárez- same genus or family can coexist if their Morales (2000). temporal and spatial occurrence, their body size, and/or their ecological requirements are Leptodiaptomus siciloides (Lilljeborg, different. In this case A. dorsalis is about 16% 1889 in De Guerne & Richard): This species smaller than M. nesus. Looking at the con- was only found in Laguna de Ayarza, a figure- ductivity of the water bodies where they were 8-shaped doublecaldera lake in the Guatemalan found, we can say that M. nesus has a higher

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 60 (1): 187-202, March 2012 199 median and a greater range of variation, but diferentes salinidades. Además, se confirmó que los diap- the outlier in A. dorsalis was as high (4 310µS/ tomidos más comunes en la Península de Yucatán fueron: Arctodiaptomus dorsalis y Mastigodiaptomus nesus. El cm Lake Salpeten, Guatemala) as in M. nesus primero se registró también en el lago de Amatitlán. Mas-

(4 340µS/cm in Loché pond, Mexico). Apart tigodiaptomus nesus está ampliamente distribuido al igual from the differences, an overlap in conductivity que A. dorsalis, pero se encuentra ausente en el área del data is given (Fig. 6). Lago Petén en Guatemala. Mastigodiaptomus reidae fue A. dorsalis also coexists with Pseudodi- localizado en dos hábitats poco profundos, éstos especí- aptomus marshi menes difieren de los tipos de la localidad, por tener un in Lago Izabal (this study, grupo de espinas de gran tamaño en la última parte de la Brinson & Nordlie 1975), Laguna de Baca- caja toráxica y los segmentos del urosoma de las hembras. lar, Cenote Mayan Blue and Puente Milagros Leptodiaptomus siciloides se encontró sólo en el Lago (Elías-Gutiérrez et al. 2008b), with Prionodi- Ayarza, el cual contiene alta salinidad. Prionodiaptomus aptomus colombiensis in L. Vicario Lagoon colombiensis se localizó en las tierras altas de Guatemala and Matillas Lagoon (Gutiérrez-Aguirre & en el Lago de Güija y en la zona del Petén en Laguna Sac- puy. Se contribuye con registros de presencia de especies Suárez-Morales 2001), and with Mastigodiap- para un mejor conocimiento de la distribución geográfica tomus texensis in Mytza Quarry (Flores-Nava de algunos copépodos calanoides. De la misma forma, los 1994). M. nesus was found together with P. hallazgos morfológicos en algunas especies son de gran marshi in Laguna de Bacalar and Almond Hill valor taxonómico para la diferenciación de especies. Lagoon, Belize (this study). Palabras clave: América Central, zooplancton, copépo- dos calanoides, Pseudodiaptomidae, Diaptomidae, ACKNOWLEDGMENTS distribución geográfica.

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