Parasegmental appendage allocation in annelids and arthropods and the homology of parapodia and arthropodia

The new animal phylogeny disrupts the traditional taxon Articulata (uniting arthropods and annelids) and thus calls into question the homology of the body segments and appendages in the two groups. Recent work in the annelid Platynereis dumerilii has shown that although the set of genes involved in body segmentation is similar in the two groups, the body units of annelids correspond to arthropod parasegments not segments. This challenges traditional ideas about the homology of "segmental" organs in annelids and arthropods, including their appendages. Here I use the expression of engrailed, wingless and Distal-less in the arthropod Artemia franciscana to identify the parasegment boundary and the appendage primordia. I show that the early body organization including the appendage primordia is parasegmental and thus identical to the annelid organization and by deriving the different adult appendages from a common ground plan I suggest that annelid and arthropod appendages are homologous structures despite their different positions in the adult animals. This also has implications for the new animal phylogeny, because it suggests that Urprotostomia was not only parasegmented but also had parasegmental appendages similar to extant annelids, and that limb-less forms in the Protostomia are derived from limb-bearing forms.     

Co-option of an anteroposterior head axis patterning system for proximodistal patterning of appendages in early bilaterian evolution

The enormous diversity of extant animal forms is a testament to the power of evolution, and much of this diversity has been achieved through the emergence of novel morphological traits. The origin of novel morphological traits is an extremely important issue in biology, and a frequent source of this novelty is co-option of pre-existing genetic systems for new purposes (Carroll et al., 2008). Appendages, such as limbs, fins and antennae, are structures common to many animal body plans which must have arisen at least once, and probably multiple times, in lineages which lacked appendages. We provide evidence that appendage proximodistal patterning genes are expressed in similar registers in the anterior embryonic neurectoderm of Drosophila melanogaster and Saccoglossus kowalevskii (a hemichordate). These results, in concert with existing expression data from a variety of other animals suggest that a pre-existing genetic system for anteroposterior head patterning was co-opted for patterning of the proximodistal axis of appendages of bilaterian animals.     

Anatomical network analysis of the musculoskeletal system reveals integration loss and parcellation boost during the fins‐to‐limbs transition

Tetrapods evolved from within the lobe‐finned fishes around 370 Ma. The evolution of limbs from lobe‐fins entailed a major reorganization of the skeletal and muscular anatomy of appendages in early tetrapods. Concurrently, a degree of similarity between pectoral and pelvic appendages also evolved. Here, we compared the anatomy of appendages in extant lobe‐finned fishes (Latimeria and Neoceratodus) and anatomically plesiomorphic amphibians (Ambystoma, Salamandra) and amniotes (Sphenodon) to trace and reconstruct the musculoskeletal changes that took place during the fins‐to‐limbs transition. We quantified the anatomy of appendages using network analysis. First, we built network models—in which nodes represent bones and muscles, and links represent their anatomical connections—and then we measured network parameters related to their anatomical integration, heterogeneity, and modularity. Our results reveal an evolutionary transition toward less integrated, more modular appendages. We interpret this transition as a diversification of muscle functions in tetrapods compared to lobe‐finned fishes. Limbs and lobe‐fins show also a greater similarity between their pectoral and pelvic appendages than ray‐fins do. These findings on extant species provide a basis for future quantitative and comprehensive reconstructions of the anatomy of limbs in early tetrapod fossils, and a way to better understand the fins‐to‐limbs transition.     

Anatomy and lifestyle of Kunmingella (Arthropoda, Bradoriida) from the Chengjiang fossil Lagerstätte (lower Cambrian; Southwest China)

An updated reconstruction of the body plan, functional anatomy and life attitude of the bradoriid arthropod Kunmingella is proposed, based on new fossil specimens with preserved soft parts found in the lower Cambrian of Chengjiang and Haikou (Yunnan, SW China) and on previous evidence. The animal has a single pair of short antennae pointing towards the front (a setal pattern indicates a possible sensory function). The following set of seven appendages (each composed of a 5-segmented endopod and a leaf-like exopod fringed with setae) is poorly differentiated, except the first three pairs (with possible rake-like endopodial outgrowths, smaller exopods) and the last pair of appendages (endopod with longer and more slender podomeres). The endopods are interpreted as walking legs with a possible role in handling food particles (marginal outgrowth with setae). The leaf-like exopods may have had a respiratory function. The trunk end is short, pointed, flanked with furcal-like rami and projects beyond the posterior margin of the carapace. The attachment of the body to the exoskeleton is probably cephalic and apparently lacks any well-developed adductor muscle system. The inferred life attitude of Kunmingella (e.g. crawling on the surface of the sediment) was that of a dorsoventrally flattened arthropod capped by a folded dorsal shield (ventral gape at least 120°), thus resembling the living ostracode Manawa. The animal was also probably able to close its carapace as a response to environmental stress or to survive unfavourable conditions (e.g. buried in sediment). The anterior lobes of the valves are likely to have accommodated visual organs (possibly lensless receptors perceiving ambient light through the translucent head shield). Preserved eggs or embryos suggest a possible ventral brood care. The presence of Kunmingella in coprolites and its numerical abundance in Chengjiang sediment indicate that bradoriids constituted an important source of food for larger predators. Kunmingella differs markedly from the representatives of the crown group Crustacea (extant and Cambrian taxa) and from the stem group derivatives of Crustacea (exemplified by phosphatocopids and some ‘Orsten’ taxa) in showing no major sign of limb specialization (e.g. related to feeding strategies). Although it resembles other Chengjiang euarthropods in important aspects of its body plan (e.g. uniramous antennae, endopod/exopod configuration), Kunmingella possesses several features (e.g. antennal morphology, post-antennular appendages with 5-segmented endopods) which support the view that bradoriids may be very early derivatives of the stem line Crustacea.     

Structure and homology of Psittacosaurus tail bristles

We examined bristle‐like appendages on the tail of the Early Cretaceous basal ceratopsian dinosaur Psittacosaurus with laser‐stimulated fluorescence imaging. Our study reveals previously unknown details of these structures and confirms their identification as integumentary appendages. For the first time, we show that most bristles appear to be arranged in bundles and that they exhibit a pulp that widens towards the bristle base. We consider it likely that the psittacosaur bristles are structurally and developmentally homologous to similar filamentous appendages of other dinosaurs, namely the basal heterodontosaurid Tianyulong and the basal therizinosauroid theropod Beipiaosaurus, and attribute the greater robustness of the bristles of Psittacosaurus to a higher degree of cornification and calcification of its integument (both skin and bristles). Although the psittacosaur bristles are probably homologous with avian feathers in their origin from discrete cell populations, it is uncertain whether they developed from a follicle, one of the developmental hallmarks of true feathers. In particular, we note a striking resemblance between the psittacosaur bristles and the cornified spine on the head of the horned screamer, Anhima cornuta, an extant anseriform bird. Similar, albeit thinner keratinous filaments of extant birds are the ‘beard’ of the turkey, Meleagris gallopavo, and the crown of the Congo peafowl, Afropavo congensis. All of these structures of extant birds are distinct from true feathers, and because at least the turkey beard does not develop from follicles, detailed future studies of their development would be invaluable towards deepening our understanding of dinosaur filamentous integumentary structures.     

A chiton without a foot

Abstract: The palaeoloricate ‘polyplacophorans’ are an extinct paraphyletic group of basal chiton‐like organisms known primarily from their fossilized valves. Their phylogenetic placement remains contentious, but they are likely to include both stem‐group Polyplacophora and stem‐group Aplacophora. Candidates for the latter position include ‘Helminthochiton’thraivensis from the Ordovician of Scotland, which we redescribe here through a combined optical and micro‐CT (XMT) restudy of the type material. The 11 specimens in the type series are all articulated, presenting partial or complete valve series as well as mouldic preservation of the girdle armature; they demonstrate a vermiform body plan. The valves are typically palaeoloricate in aspect, but differ in detail from all existing palaeoloricate genera; we hence erect Phthipodochiton gen. nov. to contain the species. The most notable feature of the fossils is the spicular girdle; this is impersistently preserved, but demonstrably wraps entirely around the ventral surface of the animal, implying that a ‘true’ (i.e. polyplacophoran like) foot was absent, although we do not exclude the possibility of a narrow solenogastre‐like median pedal groove having been present. Phthipodochiton thraivensis presents an apparent mosaic of aplacophoran and polyplacophoran features and as such will inform our understanding of the relationship between these groups of extant molluscs. An inference may also be drawn that at least some other palaeoloricates possessed an ‘armoured aplacophoran’ body plan, in contrast to the ‘limpet‐like’ body plan of extant Polyplacophora.     

Mitochondrial genome of a Japanese placozoan

Placozoans are marine invertebrates found in tropical and subtropical waters. Their body plan is among the simplest of free-living animals. The present study determined the mitochondrial genome sequence of a placozoan collected on the coast of Shirahama, Wakayama, Honshu, Japan, and compared it with those of Trichoplax adhaerens from the Red Sea and of three strains from the Caribbean Sea. The sequences of mitochondrial respiratory chain of the Japanese placozoan genes are very similar to those of the BZ49 strain from the Caribbean Sea. However, there are distinct differences in gene arrangement, such as the location of two open reading frames. This Japanese placozoan is therefore distinguishable from the other strains. Based on current knowledge of placozoan 16S diversity our 'Shirahama' strain most likely represents the H15 lineage, known from the Philippines. In the mitochondrial genome of placozoans, substitution rates are slower than in bilaterians, whereas the rate of rearrangements is faster.     

The Collins’ monster,a spinous suspension-feeding lobopodian from the Cambrian Burgess Shale of British Columbia

Lobopodians, a paraphyletic group of rare but morphologically diverse Palaeozoic vermiform animals bearing metameric appendages, are key to the origin of extant panarthropods. First discovered in 1983 on Mount Stephen (Yoho National Park, British Columbia), the Cambrian (Wuliuan) Burgess Shale lobopodian nicknamed ‘Collins’ monster’ is formally described as Collinsovermis monstruosus gen. et sp. nov. A formal systematic treatment of the comparable and poorly known lobopodian Acinocricus stichus from Utah is also provided. The body of Collinsovermis is plump and compact but shows the diagnostic suspension-feeding characters of luolishaniid lobopodians. It possesses 14 contiguous pairs of lobopods, lacking space between them. The 6 anterior pairs are elongate, adorned with about 20 pairs of long and slightly curved ventral spinules arranged in a chevron-like pattern. These appendages terminate in a pair of thin claws and their dorsal surfaces are covered in minute spines or setae. The 8 posterior lobopod pairs, which attach to a truncated body termination, are stout and smooth, each terminated by a single strong recurved claw. Each somite bears a pair of dorsal spines; somites 4 and posteriad bear an additional median spine. The spines on somites 1–3 are much shorter than the spines on the remaining somites. The head is short, bears a terminal mouth and a pair of antenniform outgrowths, and is covered by an oblong sclerite. Collinsovermis, plus Collinsium and Acinocricus, are found to comprise a sub-group of stout luolishaniid lobopodians with remarkably long spinules on the front lobopods, interpreted here as a clade (Teratopodidae fam. nov.) This clade is distinct from both the comparatively slenderer Luolishania and a sub-group composed of Facivermis and Ovatiovermis lacking body sclerites. Luolishaniids were mostly sessile forerunners of arthropods that had coupled efficient suspension-feeding devices and, as in Collinsovermis, strong defensive or deterrent features.     

Characterization of a Novel <Emphasis Type="Italic">Bacillus thuringiensis</Emphasis> Phenotype Possessing Multiple Appendages Attached to a Parasporal Body

Bacillus thuringiensis is a bacterium best known for its production of crystal-like bodies comprised of one or more Cry-proteins, which can be toxic to insects, nematodes or cancer cells. Although strains of B. thuringiensis have occasionally been observed with filamentous appendages attached to their spores, appendages in association with their parasporal bodies are extremely rare. Herein we report the characterization of Bt1-88, a bacterial strain isolated from the Caribbean that produces a spore–crystal complex containing six long appendages, each comprised of numerous thinner filaments approximately 10 nm in diameter and 2.5 μm in length. Each of the multi-filament appendages was attached to a single, small parasporal body located at one end of the bacterial spore. Biochemical tests, 16S rDNA gene sequencing, and the identification of two Cry proteins by partial protein sequencing (putatively Cry1A and Cry2A), unambiguously identified Bt1-88 as a strain of B. thuringiensis. Bt1-88 represents the second reported strain of B. thuringiensis possessing a parasporal body/appendage phenotype characterized by one or more long appendages, comprised of numerous filaments in association with a parasporal body. This finding suggests that Bt1-88 is a member of a new phenotypic class of B. thuringiensis, in which the parasporal body may perform a novel structural role through its association with multi-filament appendages.     

Cellular and molecular processes leading to embryo formation in sponges: evidences for high conservation of processes throughout animal evolution

The emergence of multicellularity is regarded as one of the major evolutionary events of life. This transition unicellularity/pluricellularity was acquired independently several times (King 2004). The acquisition of multicellularity implies the emergence of cellular cohesion and means of communication, as well as molecular mechanisms enabling the control of morphogenesis and body plan patterning. Some of these molecular tools seem to have predated the acquisition of multicellularity while others are regarded as the acquisition of specific lineages. Morphogenesis consists in the spatial migration of cells or cell layers during embryonic development, metamorphosis, asexual reproduction, growth, and regeneration, resulting in the formation and patterning of a body. In this paper, our aim is to review what is currently known concerning basal metazoans—sponges’ morphogenesis from the tissular, cellular, and molecular points of view—and what remains to elucidate. Our review attempts to show that morphogenetic processes found in sponges are as diverse and complex as those found in other animals. In true epithelial sponges (Homoscleromorpha), as well as in others, we find similar cell/layer movements, cellular shape changes involved in major morphogenetic processes such as embryogenesis or larval metamorphosis. Thus, sponges can provide information enabling us to better understand early animal evolution at the molecular level but also at the cell/cell layer level. Indeed, comparison of molecular tools will only be of value if accompanied by functional data and expression studies during morphogenetic processes.     

Origination of antlerogenesis

Antlers are unique appendages. They are shed and rebuilt at intervals, and are synapomorphic for all living Cervidae (except for the Chinese water deer, Hydropotes inermis , in which they have presumably been lost). The antlerogenic process is controlled by a complex interaction of fluctuating levels of several hormones, most importantly testosterone. The oldest antler remains are recorded from the early Miocene; these have been interpreted as non‐deciduous appendages because of supposed permanent skin coverage and the lack of a burr (a well‐developed osseous protuberance around the base of the antler, which is always present in extant cervids). The aim of this study is to test the hypothesis that antler shedding was possible in these early Miocene cervids. Antlers of all extant and eight Miocene cervid genera, including burr‐less antler fragments of the earliest cervids Procervulus , Ligeromeryx , and Lagomeryx were studied. An extensive comparative morphological analysis of external features of the antler, and of the abscission area and the base of the antler in particular, was undertaken. The results indicate that a regular, porous, and rugose abscission surface at the proximal end of the antler indicates antler shedding in both living and fossil cervids. The antler shedding mechanism must therefore have already been present in all early/mid Miocene cervid genera included in this study. On this basis, it is suggested that the presence of a burr is not prerequisite in order to shed antlers, that the presence of perpetual antlers has not yet been verified, and that the process of shedding and regeneration developed with the first appearance of these organs. This insight is particularly important for the systematic classification of early Miocene species as Cervidae, because the absence of the antler shedding and rebuilding mechanism would exclude them from the taxon Cervidae and from any relationship with extant cervids. J. Morphol. 278:182–202, 2017. © 2016 Wiley Periodicals,Inc.     

Deciphering the early evolution of echinoderms with Cambrian fossils

Echinoderms are a major group of invertebrate deuterostomes that have been an important component of marine ecosystems throughout the Phanerozoic. Their fossil record extends back to the Cambrian, when several disparate groups appear in different palaeocontinents at about the same time. Many of these early forms exhibit character combinations that differ radically from extant taxa, and thus their anatomy and phylogeny have long been controversial. Deciphering the earliest evolution of echinoderms therefore requires a detailed understanding of the morphology of Cambrian fossils, as well as the selection of an appropriate root and the identification of homologies for use in phylogenetic analysis. Based on the sister‐group relationships and ontogeny of modern species and new fossil discoveries, we now know that the first echinoderms were bilaterally symmetrical, represented in the fossil record by Ctenoimbricata and some early ctenocystoids. The next branch in echinoderm phylogeny is represented by the asymmetrical cinctans and solutes, with an echinoderm‐type ambulacral system originating in the more crownward of these groups (solutes). The first radial echinoderms are the helicoplacoids, which possess a triradial body plan with three ambulacra radiating from a lateral mouth. Helicocystoids represent the first pentaradial echinoderms and have the mouth facing upwards with five radiating recumbent ambulacra. Pentaradial echinoderms diversified rapidly from the beginning of their history, and the most significant differences between groups are recorded in the construction of the oral area and ambulacra, as well as the nature of their feeding appendages. Taken together, this provides a clear narrative of the early evolution of the echinoderm body plan.     

How the pterosaur got its wings

Throughout the evolutionary history of life, only three vertebrate lineages took to the air by acquiring a body plan suitable for powered flight: birds, bats, and pterosaurs. Because pterosaurs were the earliest vertebrate lineage capable of powered flight and included the largest volant animal in the history of the earth, understanding how they evolved their flight apparatus, the wing, is an important issue in evolutionary biology. Herein, I speculate on the potential basis of pterosaur wing evolution using recent advances in the developmental biology of flying and non‐flying vertebrates. The most significant morphological features of pterosaur wings are: (i) a disproportionately elongated fourth finger, and (ii) a wing membrane called the brachiopatagium, which stretches from the posterior surface of the arm and elongated fourth finger to the anterior surface of the leg. At limb‐forming stages of pterosaur embryos, the zone of polarizing activity (ZPA) cells, from which the fourth finger eventually differentiates, could up‐regulate, restrict, and prolong expression of 5′‐located Homeobox D (Hoxd) genes (e.g. Hoxd11, Hoxd12, and Hoxd13) around the ZPA through pterosaur‐specific exploitation of sonic hedgehog (SHH) signalling. 5′Hoxd genes could then influence downstream bone morphogenetic protein (BMP) signalling to facilitate chondrocyte proliferation in long bones. Potential expression of Fgf10 and Tbx3 in the primordium of the brachiopatagium formed posterior to the forelimb bud might also facilitate elongation of the phalanges of the fourth finger. To establish the flight‐adapted musculoskeletal morphology shared by all volant vertebrates, pterosaurs probably underwent regulatory changes in the expression of genes controlling forelimb and pectoral girdle musculoskeletal development (e.g. Tbx5), as well as certain changes in the mode of cell–cell interactions between muscular and connective tissues in the early phase of their evolution. Developmental data now accumulating for extant vertebrate taxa could be helpful in understanding the cellular and molecular mechanisms of body‐plan evolution in extinct vertebrates as well as extant vertebrates with unique morphology whose embryonic materials are hard to obtain.     

Ancestral Patterning of Tergite Formation in a Centipede Suggests Derived Mode of Trunk Segmentation in Trilobites

Trilobites have a rich and abundant fossil record, but little is known about the intrinsic mechanisms that orchestrate their body organization. To date, there is disagreement regarding the correspondence, or lack thereof, of the segmental units that constitute the trilobite trunk and their associated exoskeletal elements. The phylogenetic position of trilobites within total-group Euarthropoda, however, allows inferences about the underlying organization in these extinct taxa to be made, as some of the fundamental genetic processes for constructing the trunk segments are remarkably conserved among living arthropods. One example is the expression of the segment polarity gene engrailed, which at embryonic and early postembryonic stages is expressed in extant panarthropods (i.e. tardigrades, onychophorans, euarthropods) as transverse stripes that define the posteriormost region of each trunk segment. Due to its conservative morphology and allegedly primitive trunk tagmosis, we have utilized the centipede Strigamia maritima to study the correspondence between the expression of engrailed during late embryonic to postembryonic stages, and the development of the dorsal exoskeletal plates (i.e. tergites). The results corroborate the close correlation between the formation of the tergite borders and the dorsal expression of engrailed, and suggest that this association represents a symplesiomorphy within Euarthropoda. This correspondence between the genetic and phenetic levels enables making accurate inferences about the dorsoventral expression domains of engrailed in the trunk of exceptionally preserved trilobites and their close relatives, and is suggestive of the widespread occurrence of a distinct type of genetic segmental mismatch in these extinct arthropods. The metameric organization of the digestive tract in trilobites provides further support to this new interpretation. The wider evolutionary implications of these findings suggest the presence of a derived morphogenetic patterning mechanism responsible for the reiterated occurrence of different types of trunk dorsoventral segmental mismatch in several phylogenetically distant, extinct and extant, arthropod groups.     

The evolution of mammalian body temperature: the Cenozoic supraendothermic pulses

In this study, I investigated the source(s) of variation in the body temperatures of mammals. I also attempted to reconstruct ancestral normothermic rest-phase body temperature states using a maximum parsimony approach. Body temperature at the familial level is not correlated with body mass. For small mammals, except the Macroscelidae, previously identified correlates, such as climate adaptation and zoogeography explained some, but not all, T _b apomorphies. At the species level in large cursorial mammals, there was a significant correlation between body temperature and the ratio between metatarsal length and femur length, the proxy for stride length and cursoriality. With the exception of two primate families, all supraendothermic (T _b > 37.9°C) mammals are cursorial, including Artiodactyla, Lagomorpha, some large Rodentia, and Carnivora. The ruminant supraendothermic cursorial pulse is putatively associated with global cooling and vegetation changes following the Paleocene–Eocene Thermal Maximum. Reconstructed ancestral body temperatures were highly unrealistic deep within the mammalian phylogeny because of the lack of fossil T _b data that effectively creates ghost lineages. However, it is anticipated that the method of estimating body temperature from the abundance of ¹³C–¹⁸O bonds in the carbonate component of tooth bioapatite in both extant and extinct animals may be a very promising tool for estimating the T _b of extinct mammals. Fossil T _b data are essential for discerning derived T _b reversals from ancestral states, and verifying the dates of supraendothermic pulses.     

A comparison of swimming structures and kinematics in three species of crustacean larvae

Crustacean larvae swim with paired rowing appendages that rotate around the body of the animal. The number of paired rowing appendages varies from one species of larvae to another. In addition, the size of the crustacean larvae is different between species and increases as they grow. The nature of the fluid forces changes as size increases, so the morphology and mechanics of swimming in these animals will change during increases in size. This article demonstrates the changing kinematics of locomotion between three species of crustacean larvae, which swim with one (Artemia franciscana), two (Carcinus maenas) or five (Homarus americanus) pairs of propulsive limbs. The relative change in the surface area and volume ratios of the locomotor structures are also demonstrated.     

Saccoglossus testa from the Mazon Creek fauna (Pennsylvanian of Illinois) and the evolution of acorn worms (Enteropneusta: Hemichordata)

The limited fossil record of enteropneust hemichordates (acorn worms) and the few external features that distinguish the four families have provided a challenge to our understanding of the evolution of the group and their various feeding adaptations. The middle Pennsylvanian Saccoglossus testa sp. nov. from the Mazon Creek, Westfalian D Carbonate Formation, Francis Creek Shale of northern Illinois provides evidence for the exploitation of surface sediments. Saccoglossus testa has a long proboscis characteristic of the extant genus Saccoglossus, a specialist in surface deposit feeding. The collar is as long as it is wide. The anterior trunk lacks a distinctively wide branchial region. These three features distinguish it from its sympatric enteropneust species Mazoglossus ramsdelli Bardack that has a proboscis characteristic of an infaunal deposit feeder. It is the seventh known species of fossil enteropneust, including a resting trace of a Lower Triassic fossil that has collar lips that characterize the extant deep‐sea family Torquaratoridae, and which represents a second parallel evolution of surface deposit feeding. An analysis of the seven fossils shows that the earliest Enteropneusta had a relatively simple harrimaniid‐like body plan, and that the spengelid, the torquaratorid and lastly the most complex ptychoderid body plan appeared in that chronological order.     

Seasonal morphometric variation in sexual and asexual, North American and Australian, populations of the aphid, Acyrthosiphon pisum

The lengths of the body and appendages of the aphid Acyrthosiphon pisum (Harris) (Homoptera: Aphididae) vary seasonally in sexual North American and asexual Australian populations. The first generation of spring aphids in North America and winter aphids in Australia have short appendages in relation to body length. Excluding this phenotype, North American and Australian aphids cannot be discriminated morphometrically. The short appendages in North America are associated with a specialized morph called a fundatrix; the short appendages of Australian aphids are caused by exposure to low temperatures during prenatal development. The same temperature-sensitive mechanism operates in sexual and asexual North American aphids, but does not explain the short appendages of the fundatrix, which appear to arise through a separate mechanism. The short appendages are caused neither by a maternal effect from winged mothers, although such an effect exists, nor by seasonal changes in body length and allometry, nor by microevolutionary changes. The temperature-induced shortening of appendages is a seasonal polymorphism, which mimics the short appendages seen in fundatrices. The two types of phenotypic plasticity have the same consequence in sexual and asexual populations of the same species and may be an example of convergent evolution.