A NEW SPINICAUDATAN GENUS (CRUSTACEA: ‘CONCHOSTRACA’) FROM THE LATE CRETACEOUS OF MADAGASCAR

Abstract: A new spinicaudatan genus and species, Ethmosestheria mahajangaensis gen. et sp. nov., is described from the Anembalemba Member (Upper Cretaceous, Maastrichtian) of the Maevarano Formation, Mahajanga Basin, Madagascar. This is the first spinicaudatan reported from the post‐Triassic Mesozoic of Madagascar. The new species is assigned to the family Antronestheriidae based on the cavernous or sievelike ornamentation on the carapace. Of well‐documented Mesozoic spinicaudatan genera, Ethmosestheria mahajangaensis is most closely related to Antronestheria Chen and Hudson from the Great Estuarine Group (Jurassic) of Scotland. However, relatively poor documentation of the ornamentation of most Gondwanan Mesozoic spinicaudatan species precludes detailed comparison among taxa. Ethmosestheria mahajangaensis exhibits ontogenetic trends in carapace growth: a change in carapace outline from subcircular/subelliptical to elliptical, and from very wide juvenile growth bands to narrow adult growth bands. Ornamentation style, however, does not vary with ontogeny. Ethmosestheria mahajangaensis individuals lived in temporary pools in a broad channel‐belt system within a semiarid environment; preserved desiccation structures on carapaces indicate seasonal drying out of pools within the river system. Specimens of Ethmosestheria mahajangaensis are preserved with exquisite detail in debris flow deposits; these are the first spinicaudatans reported from debris flow deposits. These deposits also contain a varied vertebrate fauna, including dinosaurs, crocodyliforms, turtles, and frogs. Rapid entombment of the spinicaudatan carapaces likely promoted early fossil diagenesis leading to highly detailed preservation.     

Evidence for a dioecious mating system in Early Jurassic Hardapestheria maxwelli gen. et sp. nov. (Crustacea,Branchiopoda, Spinicaudata) from the Kalkrand Formation of Namibia

A new spinicaudatan (clam shrimp), Hardapestheria maxwelli gen. et sp. nov., is described from the Jurassic Kalkrand Formation of central Namibia. Specimens were collected from a sedimentary interbed within a succession of flood basalts. These are the first spinicaudatans to be described from the Jurassic of south‐western Africa. The new taxon is assigned to the family Eosestheriidae based on the combination of punctae and radial ornamentation on the carapace. Ornamentation on the growth bands in H. maxwelli differs from other eosestheriid genera because the punctate ornamentation is not restricted to the dorsal region of the carapace. Instead, all growth bands include a proximal region with punctae even though the distal portion of each growth band may exhibit anastomosing lirae. Among well‐documented Mesozoic spinicaudatan genera, Hardapestheria is most closely related to Carapacestheria from the contemporaneous Kirkpatrick Basalt of Antarctica and Yanjiestheria from the Early Cretaceous of China. Hardapestheria maxwelli displays sexual dimorphism, which can be definitively related to a dioecious mating system with discrete male and female individuals. Review of additional early members of the Eosestheriidae suggests that the maintenance of two discrete sexes was the ancestral state for the clade. The ability to discriminate sexual mating system type unambiguously is rare in fossils, and this new species sheds light on the relationship between environmental stability and mating system evolution.     

Taxonomy, morphology and speciation of the Semicytherura henryhowei group (Crustacea, Ostracoda)

The genus Semicytherura belongs to the family Cytheruridae, and was distinguished from Cytherura on the basis of carapace features. Species of Semicytherura from Japan and adjacent seas can be divided into two groups. One is represented by Semicytherura miurensis Hanai, 1957, characterized by a thin, oval carapace covered with fine reticulation. The other is represented by Semicytherura henryhowei Hanai & Ikeya, 1977, characterized by a thick sub-rectangular carapace in lateral view. Semicytherura henryhowei, which is distributed from Hokkaido to Okinawa in Japan, has been regarded as having several morphotypes distinguishable on outline and reticulation of carapace. However, as a result of detailed observations on the copulatory organ, carapace outline and distributional pattern of pore systems, remarkable differences are shown to exist between the two most frequently occurring morphotypes. In order to recognize S. henryhowei sensu stricto, the carapace of the holotype was re-examined. Consequently, neither of the two morphotypes are considered to belong to S. henryhowei due to differences of carapace outline and distribution of pore systems. The two morphotypes are here regarded as independent taxa, described as new: S. kazahanan. sp. and S. sasameyukin. sp. The geographical distributions of the two new species overlap, but their micro-habitats differ from each other; the former lives on calcareous algae on rocky shores, the latter lives on silty sand bottom within the inner bay. A third new species, S. slipperi sp. nov., is also described. In view of their present geographical distributions and fossil records, the origin of this group of species would appear to be the Japanese islands or adjacent areas in and after the Miocene. This group then migrated to the Arctic Ocean and East Pacific Ocean during or before the middle Pliocene.     

Thylacocephala (Arthropoda: Crustacea?) from the Cretaceous of Lebanon and implications for thylacocephalan systematics

An intensive study of a collection of arthropods from the Cretaceous of Lebanon, formerly referred to as stomatopod larvae, reveals that these forms belong to the problematical arthropod class Thylacocephala. The species Protozoea hilgendorfi , P. damesi , and Pseuderichthus cretaceus display defining thylacocephalan characters such as a carapace enclosing the entire body bearing a large anterior optic notch; three pairs of large, raptorial appendages; and a posterior battery of small swimming limbs associated with muscle segments. Unique characters of at least the genus Protozoea are the numerous 'pits' covering the entire carapace, elongated anterior rostral and posterior spines, and a dorsal hinge indicative of a truly bivalved carapace. We note several anatomical curiosities that provide some insight into the anatomy, ecology and phylogenetic relationships of these and other thylacocephalan species. Structures on the carapace surface of thylacocephalans may represent sensory organs comparable to the receptors seen on thecostracan crustaceans. The affinities of the three species described are still not entirely clear, but a phylogenetic analysis of the entire group may resolve this.     

A new genus and species of raninoid crab from the Upper Cretaceous of Mississippi

A new genus and species of raninoid crab, Carinatus galebishopi, is described from the upper Maastrichtian Prairie Bluff and Owl Creek formations of Mississippi. Carinatus galebishopi gen. et sp. nov. fits best within the Raninidae, Subfamily Symethinae, based upon similar dorsal carapace characteristics of member genera. The new crab possesses distinctive dorsal carapace ornamentation setting it apart from described symethines and warrants establishment of a new genus. Carinatus galebishopi represents the first Cretaceous symethine crab and suggests the subfamily may have migrated eastward across the North Atlantic—Maastrichtian of Mississippi, Danian of New Jersey, and Ypresian of Spain—before dispersing to the Indo-Pacific.     

The largest Cretaceous podocnemidoid turtle (Pleurodira) revealed by an isolated plate from the Bauru Basin,south-central Brazil

The Bauru Basin (south-central Brazil) fossils have largely contributed to understand the faunal composition of South American Cretaceous. Among those, several turtle specimens were retrieved from those deposits, all belonging to Podocnemidoidae, the single group known from the Basin. On the other hand, only incomplete shell elements indicate large turtles such as ‘Peirópolis A’. Another shell fragment, a large peripheral plate from the Marília Formation, is described here. Its lack of surface ornamentation, and deep sulci are generally typical for podocnemidoids. The plate exhibits a narrow knob slightly projected onto the pleuro-marginal sulcus, absent in any other Bauru Basin turtle. According to extant and fossil podocnemidoids measures, we estimated this individual reached 1 m of carapace length, exceeding in size all other taxa known from the Basin. This new report reveals a broader size variation among podocnemidoids from the Late Cretaceous of Bauru Basin and a morphological diversity previously unknown for the period, comparable to those of the Amazon Basin today, in which inhabits the largest extant podocnemidoid Podocnemis expansa. Furthermore, the occurrence of such large turtles implies the presence of perennial streams on the northeastern portion of the Bauru Basin during the Maastrichtian and a richer environment than previously thought.     

The structure of the ostracode carapace

The carapace ultrastructure of three Recent [Heterocypris incongruens (Ramdohr), Cypridopsis uidua (Müller), and Conchoecia belgica Müller] and one Cretaceous ostracode species [Cypridea propunctata Sylvester-Bradley] has been studied by means of standard electron microscope techniques. The carapace is shown to be an organic structure divisible into an outer epicuticle, a median exocuticle and an inner endocuticle in the three cypridacean ostracodes examined. In Conchoecia belgica only an outer epicuticle and an inner endocuticle were observed, but this may be due to preservation. The chitin structure of the carapace of calcareous ostracodes is shown to be essentially an interlocking lattice, more coarsely developed in the exocuticle; layered (lamellar) chitin is only present in the selvage and in the connective tissue joining the valves at the hinge. The non-calcareous Conchoecia belgica possesses the layered chitin structure common to other crustaceans and to insects. The relationship of the carapace structure to that present in the decapod and insect exoskeleton is discussed. The technical problems met with during this study are considered in order to outline the difficulties associated with the study of organisms of this size.     

Effects of the evolution of the Serra do Mar mountains on the shape of the geographically isolated populations of Aegla schmitti Hobbs III, 1979 (Decapoda: Anomura)

The Aeglidae family is a useful model for evaluating the shape variation to present small geographically isolated populations and present a carapace structure that allows precise placement of landmarks. We analysed variations of the size and shape of the carapace of seven populations of A. schmitti. We used 18 bidimensional anatomical landmarks on the carapace of adult males. Size variation between the populations was analysed through a univariate analysis of variance, while the shape variation was analysed through a multivariate analysis of variance using the configurations aligned by the generalized Procrustes analysis. The ordination of the populations was investigated through a canonical variables analysis. Individuals from the populations differed in the carapace size, in this case, we observed the largest individuals in the Iguaçu river basin, followed by Ribeira do Iguape and the Coastal basin. Carapace shape also differed between the populations of A. schmitti, with the exception of the Mato Grande and Piraquara rivers: populations from the Iguaçu River presented a larger and rounder carapace when compared with the slenderer carapace of the Coastal basin, and the Ribeira do Iguape animals presented intermediate shapes. Data from the present study show great congruency with the draining pattern and geological history of the region.     

Invasive Australian crayfish Cherax quadricarinatus in the Sanyati Basin of Lake Kariba: a preliminary survey

The invasion of Cherax quadricarinatus, the Australian redclaw crayfish, in the Sanyati Basin of Lake Kariba, Zimbabwe, is reported. A total of 79 crayfish were caught at 10 out of 12 sampling sites in the Sanyati Basin in November–December 2012. The average catch per unit effort (CPUE) varied from 1.1 to 4 crayfish per trap per night, carapace length ranged from 29 to 93.5 mm, and weight ranged from 4 to 196.2 g. Most crayfish were between 50 and 59.9 mm carapace length. Males (average 82.6 g) were significantly heavier than females (37.2 g) and males were larger in carapace length, carapace width, chela length and chela width. A feral population of C. quadricarinatus is now established in the Sanyati Basin. Possible modes of dispersal within the basin are discussed.     

Geometric morphometrics of carapace of Macrobrachium australe (Crustacea: Palaemonidae) from Reunion Island

Zimmermann, G., Bosc, P., Valade, P., Cornette, R., Améziane, N. and Debat, V. 2011. Geometric morphometrics of carapace of Macrobrachium australe (Crustacea: Palaemonidae) from Reunion Island. —Acta Zoologica (Stockholm) 93 : 492–500. We investigated the structure of carapace shape variation in six populations of Macrobrachium australe Guérin‐Méneville 1838 (Crustacea: Decapoda: Palaemonidae) from Reunion Island (Indian Ocean) freshwaters. The morphometric analysis revealed the occurrence of two morphotypes corresponding to two different types of habitats. Individuals living in lotic habitats present a thick carapace armed with a short, robust and straight rostrum, while individuals from lentic habitats have a slender carapace armed with a thin long rostrum orientated upward. This difference suggests an adaptation to lotic disturbances and is tentatively interpreted as adaptive phenotypic plasticity. In such amphidromous organisms regressing to freshwaters after a marine larval phase, selection for physiological and developmental flexibility might facilitate further adaptation and allows the colonisation of a wide panel of environmentally different and sometimes geographically distant insular streams.     

First Jurassic occurence of Enoploclytia M’Coy, 1849 (Crustacea: Decapoda: Erymidae)

Among the Erymidae Van Straelen, 1925, known as early as the Late Permian (Changhsingian) and widespread in the Jurassic, the genus Enoploclytia M’Coy, 1849 seems to have a late appearance because of its lack before the Cretaceous. Until now, the oldest representative was E. augustobonae Devillez, Charbonnier, Hy?ný and Leroy, 2016 from the Early Cretaceous (Barremian) of the eastern Paris basin (France). However, a new fossil collected in Normandy (France), showing the typical carapace groove pattern of Enoploclytia, attests the presence of the genus in the Late Jurassic (Oxfordian).     

Development of the turtle carapace: Implications for the evolution of a novel bauplan

The chelonian carapace is composed of the endochondral ribs and vertebrae associated with a specialized dermis. The ribs are found in an aberrant position compared to those of all other tetrapods; they are superficial and dorsal to the limb girdles. This morphological arrangement, which constitutes the unique chelonian Bauplan, is examined from a developmental perspective. Embryos of Chelydra serpentina were studied during stages of carapace development. Tissue morphology, autoradiography, and indirect immunofluorescent localization of adhesion molecules indicate that the outgrowth of the embryonic carapace occurs as the result of an epithelial–mesenchymal interaction in the body wall. A carapacial ridge composed of mesenchyme of the dermis and overlying ectoderm is formed dorsal to the ectodermal boundary between somitic and lateral plate mesoderm. It is the anlage of the carapace margin, in which the ribs will eventually terminate. The ectoderm of the carapacial ridge is thickened into a pseudostratified columnar epithelium, which overlies a condensation in the mesenchyme of the dermis. Patterns of cell proliferation and the distribution of N-CAM and fibronectin in the carapacial ridge are consistent with patterns seen in other structures initiated by epithelial–mesenchymal interactions such as feathers and limb buds. Based on an analogy to this developmental mechanism in the development of the limb skeleton, a further analogy with the evolution of the limbs from lateral fin folds is used to form a hypothesis on the evolution of the carapace from elements of the primitive reptilian integument.     

A first record of anchialine crab of the genus Orcovita Ng and Tomascik, 1994 (Decapoda: Brachyura: Varunidae) from Japan, with description of the species

Orcovita miruku, new species, is described from Ishigaki Island, southern Ryukyu Islands, Japan. Orcovita miruku is easily distinguished from the other congeners by the presence of three teeth (including external orbital angle) on the anterolateral margin of the carapace. Orcovita angulata may be allied to O. miruku, but there are a number of differences in the characters of the carapace and the ambulatory legs. The holotype, obtained from an anchiline pool in a limestone excavation, is the only specimen collected thus far, and this is the first record of an anchiline crab from Japan.     

New early Jurassic decapod crustacean from Patagonia (Chubut province), Argentina

A well-preserved decapod specimen was found in early Toarcian deposits cropping out on the western slope of Meseta Catreleo, central Chubut province, Argentina. It is a nearly complete exoskeleton preserved in lateral view, slightly crushed, in fine-grained sandstones. The skeleton is mostly articulated, though some pieces are disarticulated or missing. Taphonomic features indicate a relatively rapid burial after death, with little or no transport. This specimen is here described as a new species of the genus Mecochirus Germar (Decapoda: Glypheoidea). Mecochirus robbianoi n. sp. is characterized by a very long, achelate, first pair of pereiopods with a narrow, long, straight dactylus, a thin carapace ornamented by low tubercules, with a slightly oblique cervical groove and a short acute rostrum. The family Mecochiridae arose in the Triassic, but the group diversified during the Early Jurassic. Mecochirus had a wide geographical distribution and is known for sure from Lower Jurassic to Upper Cretaceous deposits. The new record from Chubut is one of the oldest for the genus, the oldest for South America, and the first for Argentina, thus considerably extending the known distribution of the genus during the Early Jurassic.     

Testing the success of palaeontological methods in the delimitation of clam shrimp (Crustacea,Branchiopoda) on extant species

Fossil spinicaudatan taxonomy heavily relies on carapace features (size, shape, ornamentation) and palaeontologists have greatly refined methods to study and describe carapace variability. Whether carapace features alone are sufficient for distinguishing between species of a single genus has remained untested. In our study, we tested common palaeontological methods on 481 individuals of the extant Australian genus Ozestheria that have been previously assigned to ten species based on genetic analysis. All species are morphologically distinct based on geometric morphometrics (p ≤ 0.001), but they occupy overlapping regions in Ozestheria morphospace. Linear discriminant analysis of Fourier shape coefficients reaches a mean model performance of 93.8% correctly classified individuals over all possible 45 pairwise species comparisons. This can be further increased by combining the size and shape datasets. Nine of the ten examined species are clearly sexually dimorphic but male and female morphologies strongly overlap within species with little influence on model performance. Ornamentation is commonly species-diagnostic; seven ornamentation types are distinguished of which six are species-specific while one is shared by four species. A transformation of main ornamental features (e.g. from punctate to smooth) can occur among closely related species suggesting short evolutionary timescales. Our overall results support the taxonomic value of carapace features, which should also receive greater attention in the taxonomy of extant species. The extensive variation in carapace shape and ornamentation is noteworthy and several species would probably have been assigned to different genera or families if these had been fossils, bearing implications for the systematics of fossil Spinicaudata.     

Turtle Shell Impression in a Coprolite from South Carolina,USA

Coprolites (fossilized feces) can preserve a wide range of biogenic components. A mold of a hatchling turtle partial shell (carapace) referable to Taphrosphys sulcatus is here identified within a coprolite from Clapp Creek in Kingstree, Williamsburg County, South Carolina, USA. The specimen is the first-known coprolite to preserve a vertebrate body impression. The small size of the turtle shell coupled with the fact that it shows signs of breakage indicates that the turtle was ingested and that the impression was made while the feces were still within the body of the predator. The detailed impression could only have survived the act of defecation if the section of bony carapace was voided concurrently and remained bonded with the feces until the latter lithified. Exceptionally, the surface texture of the scutes is preserved, including its finely pitted embryonic texture and a narrow perimeter of hatchling scute texture. The very small size of the shell represented by the impression makes it a suitable size for swallowing by any one of several large predators known from this locality. The coprolite was collected from a lag deposit containing a temporally mixed vertebrate assemblage (Cretaceous, Paleocene and Plio-Pleistocene). The genus Taphrosphys is known from both sides of the Cretaceous–Paleogene (K–Pg) boundary so, based on the size of the coprolite and the locally-known predators, the juvenile turtle could have been ingested by a mosasaur, a crocodylian, or a theropod dinosaur. Unlike mosasaurs and theropod dinosaurs, crocodylian stomachs have extremely high acid content that almost always dissolves bone. Therefore, the likely predator of this turtle was a mosasaur or a (non-avian or avian) theropod dinosaur.     

Carapace epithelia are rich in large filamentous actin bundles in Daphnia magna,Daphnia pulex,and Sida crystallina (Crustacea: Cladocera)

Cladocerans (water fleas) are planktonic crustaceans that typically have a bivalved carapace. Each valve of the carapace consists of two cuticle‐secreting epithelial layers that are separated by a hemolymphatic chamber and joined by pillar structures. Ultrastructural analyses in several species of Cladocera have shown that the carapace epithelia and pillars contain filamentous structures of unknown composition. In the present study we used a fluorescent phalloidin conjugate to show that the carapaces of three cladocerans, Daphnia magna, D. pulex, and Sida crystallina, are rich in large bundles of filamentous actin (F‐actin). In D. magna we employed confocal microscopy and orthogonal views of three‐dimensional reconstructions to show that these bundles extend radially from foci in the pillars towards the integument surfaces, and their structure is consistent with that of contractile stress fibers. Using a fluorescent lipophilic stain, DiOC₆(3), we show that the F‐actin bundles are distributed in membrane‐rich regions within the carapace epithelia, and that, in the superficial epithelium, these may be large membrane‐bound organelles. In D. magna, the F‐actin bundles are present in embryonic, juvenile instar, and adult, developmental stages, and through development the bundles become larger, contain more F‐actin, and become more widely spaced. We present an alignment of the deduced amino acid sequences of six putative D. pulex actin genes, and discuss the implications that their respective sequences have on the likelihood of their inclusion into the F‐actin bundles of the carapace. Our identification of these large F‐actin bundles within the pillars of three cladocerans provides new insight into the role these structures play in influencing carapace dynamics within this order.     

Aspects of the biology of the big-handed crab,Heterozius rotundifrons (Decapoda: Brachyura), from Kaikoura,New Zealand

Abstract

From January 1975 to June 1976, samples of Heterozius rotundifrons A. Milne Edwards, 1867 were taken monthly from the intertidal zone at First Bay, Kaikoura Peninsula (42°25′S, 173°42′E). Small crabs (< 8 mm carapace width) were never common; they were either too cryptic to be collected in quantity, or have a different habitat from larger crabs. Ovigerous females were present in all months except February (in both 1975 and 1976), but generally formed less than 50% of the total sample of females each month. The number of newly deposited eggs (0.75×0.81 mm) carried by females increased with increase in carapace width according to the equation y = -1123.56+102.97x (r ² = 0.8213). Egg development lasted 3–5 months, and egg mortality during this period was almost 10%. The overall sex ratio for the 18-month sampling period was 1846 ♀ : 993 ♂, which suggests that females were approximately twice as numerous as males. However, when crabs were sorted into size classes it was evident that the sex ratio was not significantly different from 1:1 in crabs of 6–12 mm carapace width, whereas females significantly (P<0.001) outnumbered males in the 13–21 mm size range. The right and left chela were approximately equal in length in females of all sizes, but the right chela of large males was greatly enlarged compared with the left, and with the chelae of females of comparable carapace width. Increase in the growth rate of the males’ right chela commenced at a carapace width of approximately 11.0 mm. Since the smallest ovigerous female collected also had a carapace width of 11.0 mm, it is concluded that both males and females attain sexual maturity at this size.     

Ecological stasis in Spinicaudata (Crustacea,Branchiopoda)? Early Cretaceous clam shrimp of the Yixian Formation of north‐east China occupied a broader realized ecological niche than extant members of the group

The palaeoecology of Spinicaudata, the dominant group of benthic invertebrates in many pre‐Cenozoic freshwater environments, remains poorly understood. In analogy with extant taxa, it has been oversimplified and often reduced to shallow, temporary environments characterized by few trophic levels, implying ecological stasis from the Devonian to the Recent. We excavated 43 horizons of the Lower Cretaceous Yixian Formation (Anjiagou and Hengdaozi beds) to evaluate whether spinicaudatan ecology can be simplified to such an extent. Sedimentological evidence suggests general perennial conditions during the excavated lake interval. Based on 33 226 specimen counts, we identified three arthropod‐dominated macrobenthic associations and two assemblages. Response curves indicate that the spinicaudatan Eosestheria middendorfii was exceptionally tolerant to environmental gradients, followed, in decreasing order, by mayfly larvae, water boatmen and oligochaetes. Many spinicaudatan‐yielding layers represent death after reproductive senescence rather than mass‐mortality events. Spinicaudatan size varies significantly according to faunal association. A forward stepwise regression model suggests that growth responded to population density and diversity: lower densities and higher diversities triggered fast growth, and vice versa. The proposed strong density effect on carapace size has been corroborated by rearing experiments: as for E. middendorfii, natural log regression provided the best fit for the two extant species Eulimnadia texana and Eocyzicus argillaquus. Hence, E. middendorfii was a tolerant and morphologically variable species adapted to perennial waters and the frequent presence of higher trophic levels. Although there are rare records of extant taxa co‐occurring with fish, the described Early Cretaceous environment is commonly not inhabited by extant Spinicaudata.     

Nest defense and aggressive interactions between a small benthic fish (the johnny darter Etheostoma nigrum) and crayfish

Synopsis Aggression by nest-guarding male johnny darters, Etheostoma nigrum, against intruding crayfish was investigated in laboratoy experiments and field observations. In the laboratory, darter success in chasing crayfish, Orconectes rusticus, from the nest site was inversely related to crayfish size. Small crayfish (less than 15 mm carapace length) were routinely evicted from the nest area by nips directed at the posterior end of the abdomen. Although such aggressive behavior was less successful against larger crayfish, even the largest crayfish tested (carapace length 30–32 mm) were chased from the nest area in 33% of the trials. Those large crayfish that entered nests often remained despite repeated attacks by the male johnny darter and egg predation was observed. In a small Ohio stream, openings to johnny darter nests were generally between 7 and 13 mm. Thus crayfish with a carapace height greater than 13 mm (corresponding to a carapace length greater than 29 mm) would not be able to enter johnny darter nests. In field observations, male Johnny darters successfully defended nests against another crayfish species (Orconectes sanborni, carapace length 12–29 mm). Together, aggressive behavior and small size of nest entrances allow the johnny darter to successfully reproduce in areas with abundant crayfish. The Unit is sponsored jointly by the United States Fish and Wildlife Service, The Ohio Department of NaturalResources, and The Ohio State University.