Stephen C. Weeks

Global diversity of large branchiopods (Crustacea: Branchiopoda) in freshwater

With about 500 known species worldwide, the large brachiopods are a relatively small group of primitive crustaceans. With few exceptions they live in temporary aquatic systems that are most abundant in arid and semi arid areas. As many regions remain unexplored and as especially the number of species in clam shrimps and tadpole shrimps is underestimated due to difficult identification, the species list will increase with future surveys. The Branchiopoda are monophyletic, but inter-ordinal relationships, as well as many evolutionary relationships at lower taxonomic levels are still unclear. Ongoing molecular studies will more accurately depict species diversity and phylogenetic patterns. With the exception of some anostracan families, most families are not restricted to the northern or southern hemisphere or specific zoogeographical regions. Large branchiopods are used for the assessment of the quality and function of temporary wetlands. Due to the reduction in number and quality of temporary wetlands, several species became endangered and are red listed by the IUCN.

Notes on the life history of the clam shrimp, Eulimnadia texana

Several life history measures (growth rate, egg production, molt frequency, age at maturity and lifespan) were measured on several clam shrimp hermaphrodites (Eulimnadia texana Packard) grown in a laboratory setting under optimal growth conditions. Growth rates were high early in life, and then dropped dramatically when egg production began (day 5–6). Early egg production was low, and increased until approximately day 7, after which production leveled off for several days. Reproductive senescence was noted after day 17, with clutch sizes continuously dropping until death. Average molts per day was approximately 1.1, and molting seemed to be more closely associated with egg production than with growth. Growth and egg production were negatively correlated, indicating a possible trade-off between these two traits. No other trade-offs were detected. These shrimp show typical early-colonist life history traits, displaying high initial growth, early reproduction at a high rate, and then early senescence and death.

MAINTENANCE OF ANDRODIOECY IN THE FRESHWATER SHRIMP, EULIMNADIA TEXANA: ESTIMATES OF INBREEDING DEPRESSION IN TWO POPULATIONS

Abstract.— Androdioecy is an uncommon form of reproduction in which males coexist with hermaphrodites. Androdioecy is thought to be difficult to evolve in species that regularly inbreed. The freshwater shrimp Eulimnadia texana has recently been described as both androdioecious and highly selfing and is thus anomalous. Inbreeding depression is one factor that may maintain males in these populations. Here we examine the extent of “late” inbreeding depression (after sexual maturity) in these clam shrimp using two tests: (1) comparing the fitness of shrimp varying in their levels of individual heterozygosity from two natural populations that differ in overall genetic diversity; and (2) specifically outcrossing and selfing shrimp from these same populations and comparing fitness of the resulting offspring. The effects of inbreeding differed within each population. In the more genetically diverse population, fecundity, size, and mortality were significantly reduced in inbred shrimp. In the less genetically diverse population, none of the fitness measures was significantly lowered in selfed shrimp. Combining estimates of early inbreeding depression from a previous study with current estimates of late inbreeding depression suggests that inbreeding depression is substantial (δ= 0.68) in the more diverse population and somewhat lower (δ= 0.50) in the less diverse population. However, given that males have higher mortality rates than hermaphrodites, neither estimate of inbreeding depression is large enough to account for the maintenance of males in either population by inbreeding depression alone. Thus, the stability of androdioecy in this system is likely only if hermaphrodites are unable to self‐fertilize many of their own eggs when not mated to a male or if male mating success is generally high (or at least high when males are rare). Patterns of fitness responses in the two populations were consistent with the hypothesis that inbreeding depression is caused by partially recessive deleterious alleles, although a formal test of this hypothesis still needs to be conducted.

Ancient androdioecy in the freshwater crustacean Eulimnadia

Among the variety of reproductive mechanisms exhibited by living systems, one permutation—androdioecy (mixtures of males and hermaphrodites)—is distinguished by its rarity. Models of mating system evolution predict that androdioecy should be a brief stage between hermaphroditism and dioecy (separate males and females), or vice versa. Herein we report evidence of widespread and ancient androdioecy in crustaceans in the genus Eulimnadia, based on observations of over 33 000 shrimp from 36 locations from every continent except Antarctica. Using phylogenetic, biogeographical and palaeontological evidence, we infer that androdioecy in Eulimnadia has persisted for 24–180 million years and has been maintained through multiple speciation events. These results suggest that androdioecy is a highly successful aspect of the life history of these freshwater crustaceans, and has persisted for orders of magnitude longer than predicted by current models of this rare breeding system.

Evolutionary transitions among dioecy, androdioecy and hermaphroditism in limnadiid clam shrimp (Branchiopoda: Spinicaudata)

Examinations of breeding system transitions have primarily concentrated on the transition from hermaphroditism to dioecy, likely because of the preponderance of this transition within flowering plants. Fewer studies have considered the reverse transition: dioecy to hermaphroditism. A fruitful approach to studying this latter transition can be sought by studying clades in which transitions between dioecy and hermaphroditism have occurred multiple times. Freshwater crustaceans in the family Limnadiidae comprise dioecious, hermaphroditic and androdioecious (males + hermaphrodites) species, and thus this family represents an excellent model system for the assessment of the evolutionary transitions between these related breeding systems. Herein we report a phylogenetic assessment of breeding system transitions within the family using a total evidence comparative approach. We find that dioecy is the ancestral breeding system for the Limnadiidae and that a minimum of two independent transitions from dioecy to hermaphroditism occurred within this family, leading to (1) a Holarctic, all‐hermaphrodite species, Limnadia lenticularis and (2) mixtures of hermaphrodites and males in the genus Eulimnadia. Both hermaphroditic derivatives are essentially females with only a small amount of energy allocated to male function. Within Eulimnadia, we find several all‐hermaphrodite populations/species that have been independently derived at least twice from androdioecious progenitors within this genus. We discuss two adaptive (based on the notion of ‘reproductive assurance’) and one nonadaptive explanations for the derivation of all‐hermaphroditism from androdioecy. We propose that L. lenticularis likely represents an all‐hermaphrodite species that was derived from an androdioecious ancestor, much like the all‐hermaphrodite populations derived from androdioecy currently observed within the Eulimnadia. Finally, we note that the proposed hypotheses for the dioecy to hermaphroditism transition are unable to explain the derivation of a fully functional, outcrossing hermaphroditic species from a dioecious progenitor.

The role of androdioecy and gynodioecy in mediating evolutionary transitions between dioecy and hermaphroditism in the animalia.

Dioecy (gonochorism) is dominant within the Animalia, although a recent review suggests hermaphroditism is also common. Evolutionary transitions from dioecy to hermaphroditism (or vice versa) have occurred frequently in animals, but few studies suggest the advantage of such transitions. In particular, few studies assess how hermaphroditism evolves from dioecy or whether androdioecy or gynodioecy should be an “intermediate” stage, as noted in plants. Herein, these transitions are assessed by documenting the numbers of androdioecious and gynodioecious animals and inferring their ancestral reproductive mode. Both systems are rare, but androdioecy was an order of magnitude more common than gynodioecy. Transitions from dioecious ancestors were commonly to androdioecy rather than gynodioecy. Hermaphrodites evolving from sexually dimorphic dioecious ancestors appear to be constrained to those with female‐biased sex allocation; such hermaphrodites replace females to coexist with males. Hermaphrodites evolving from sexually monomorphic dioecious ancestors were not similarly constrained. Species transitioning from hermaphroditic ancestors were more commonly androdioecious than gynodioecious, contrasting with similar transitions in plants. In animals, such transitions were associated with size specialization between the sexes, whereas in plants these transitions were to avoid inbreeding depression. Further research should frame these reproductive transitions in a theoretical context, similar to botanical studies.

The effects of pond duration on the life history traits of an ephemeral pond crustacean, Eulimnadia texana

We examined the relationship between pond duration and life history characters of the clam shrimp Eulimnadia texana, a species inhabiting ephemeral ponds in southwestern North America. Since the shrimp live in temporary habitats, we predicted that there should be high selection pressure on life history characteristics associated with rapid development (e.g., fast growth, early maturity, etc.), rather than selection for increased longevity. Pond duration was estimated using a combination of average monthly rainfall and pond size (surface to volume ratio). Shrimp that live in smaller ponds (high surface to volume ratio) in areas with low average rainfall should, on average, experience a shorter total time available for development than those in larger ponds or in areas of higher rainfall. These shrimp should have an earlier age at maturity, reduced longevity, lower fecundity, and faster growth. Five replicate populations of clam shrimp were collected as cysts from five ponds. These shrimp were raised in a common garden experiment in the laboratory. Daily measurements of growth and egg production were taken and ages at maturity and death were recorded. Shrimp from areas with higher average rainfall had slower growth, higher fecundity, greater longevity, and an earlier age at maturity than those from areas with lower average rainfall. If average rainfall is an accurate measure of pond duration, then the first three of these life history traits differ in the directions expected. However, age at maturity varied in a manner opposite to that expected, being earlier in the ponds with longer duration. Surface to volume ratio was not helpful in further resolving differences in these life history characters.

Genotypic and environmental components of variation in growth and reproduction of fish hemiclones (Poeciliopsis: Poeciliidae)

The frozen‐niche‐variation model was proposed to account for the coexistence of genetically related clones in naturally occurring unisexual populations. This model is based on two assumptions: 1) ecologically different clones have multiple independent origins from sexual ancestors; and 2) the population of sexual ancestors contains genetic variability for ecologically relevant traits. To test these assumptions, we produced 14 new “hemiclones” (nonrecombining haploid genotypes) of fish (Poeciliopsis: Poeciliidae). Our ability to synthesize many new hemiclones demonstrates the feasibility of multiple independent origins of nonrecombining genotypes. A substantial proportion (10–50%) of the phenotypic variation among hemiclones in size at birth, juvenile growth rate, and fecundity had a genetic basis. Thus, we conclude that multiple origins can give rise to an assemblage of genetically distinct hemiclones, each with a unique combination of life‐history traits. Additionally, a comparative analysis of two natural hemiclones revealed that the synthetic strains represent a broad field of variation from which natural hemiclones can be selected.

REVISION OF THE EXTANT GENERA OF LIMNADIIDAE (BRANCHIOPODA: SPINICAUDATA)

ABSTRACT The extant genera of the spinicaudatan clam shrimp family Limnadiidae are revised using morphological criteria built on previously published molecular analyses. The combined analyses demonstrate the presence of eight well defined genera, two of which are new to science and one (Paralimnadia) that is resurrected. We present the description of the new genus Afrolimnadia and the new genus and species Calalimnadia mahei n. sp. described from Mauritius Island. Both molecular and morphological data strongly support eight genera: Afrolimnadia n. gen., Calalimnadia n. gen., Eulimnadia, Imnadia, Limnadia, Limnadopsis, Metalimnadia and Paralimnadia.

Relative fitness of two hermaphroditic mating types in the androdioecious clam shrimp, Eulimnadia texana

Androdioecy (populations of males and hermaphrodites) is a rare reproductive form, being described from only a handful of plants and animals. One of these is the shrimp Eulimnadia texana, which has populations comprised of three mating types: two hermaphroditic types (monogenics and amphigenics) and males. In a recent study, the amphigenic hermaphrodites were found to be in greater abundance than that predicted from a model of this mating system. Herein, we compare the relative fitness of offspring from amphigenic and monogenic siblings, attempting to understand the greater relative abundance of the former. Populations started with offspring from selfed monogenic hermaphrodites had a net reproductive rate (R) 87% that of offspring from their amphigenic siblings. Additionally, within populations of amphigenic offspring (which included males, monogenics and amphigenics), amphigenics survived longer than monogenics. These differences help to explain the increased relative abundance of amphigenics in natural populations, but amphigenics continue to be more abundant than expected.