6-8, 10, 12: Ecdysozoa (inc. Arthropoda, Insecta) + Lophotrochozoa Flashcards

1
Q

Describe the Priapulida in terms of their basic features and taxonomic position

A

Priapulida = “Penis Worms,” marine worms, most are burrowers which capture prey by waiting upright at the surface of mud or sand

Their introvert/trunk can retract using muscles, and usually ends with curved spines (scalids) around the mouth which capture food and then retract

Simple worm-like anatomy - circular nerve ring around gut, whole body covered by chitinous cuticle

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2
Q

Describe the Nematoda in terms of their basic features and taxonomic position

A

Ecdysozoans -> Nematoia (alongside Nematomorpha)

Pseudocoelomate, parasitic “roundworms” - all environments and types of animals - cause diseases eg Ascariasis, Elephantiasis and Trichinosis

Circular nerve ring around gut, specialised “oral stylets” i.e. mouthparts (some for herbivory, some for predation) - retracted piercing device

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3
Q

Name three Nematode-caused diseases and the associated species

A

ASCARIASIS (Ascaris lumbricoides) -> spread via faeces by contaminating soil, causing pneumonia, abdominal symptoms and occasionally intestinal blockage

TRICHINOSIS - caused by Trichinella, adult worms bury into small intestines, then juveniles enter blood vessels, then skeletal muscle cells. Can be lethal but drugs are available

ELEPHANTIASIS - caused by filarial worms e.g. Wuchereria or Brugia; inflammation and lymphatic obstruction, swelling, often spread by mosquitos

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4
Q

Name the groups contained within PANARTHROPODA and the defining characteristics from other ecdysozoans

A

Tardigrada, Onychophora (Velvet Worms) and Arthropoda

All have paired walking appendages with claws and show segmentation (though difficult to find a “good” synapomorph)

*share some patterns of gene expression and brain structure, but also some key differences and some features only shared between 2 of the three phyla

*Tardigrada slightly debatable - show some features of Cycloneuralia

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5
Q

Describe the Onychophora in terms of their basic features and ecology

A

VELVET WORMS - They appear like “worms with chunky, fleshy, lobed legs,”

Have a ventral nerve chord - NOT circular around the gut

They use slimy webs to trap prey (e.g., cockroaches) then eat using mandibles and mouth

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6
Q

Describe the Tardigrada in terms of their basic features and ecology

A

“Water bears/Moss Piglets” -aquatic but most live on land, often in thin water layer on moss

Have 8 non-jointed appendages, limited circulatory and respiratory systems, complex worm-like nervous system

They have a piercing stylet mouth and sucking pharynx, evolved either for predation (e.g., of nematodes and rotifers) or for herbivory

Once thought to show major HGT with bacteria - mostly just contamination

Can undergo prolonged Cryptobiosis in response to stress (anoxybiosis, anhydrobiosis, crybiosis, etc.) and form Tuns - small sacs, evolved to survive harsh conditions

BONUS - unlike trehalose used by nematodes etc. in dehydration, Tardigrades use unique intrinsically disordered proteins (TDPs) which become rigid when dessicated

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7
Q

Describe the general current outlook on Ecdysozoan phylogeny

A

Monophyly of Ecdysozoa now generally accepted, despite prior dissent from morphologists

Difficult to resolve internal phylogeny due to Long Branches of Nematoda and Tardigrada, and limited developmental/morphological data for others

Decent evidence for Panarthropoda as a clade, but other placements within Ecdysozoa still largely debated - microRNAs may soon prove useful in resolving this due to their deep homology

Current Leading Hypothesis:

  • Scalidiphora (Priapulida, Kinorhyncha and Loricifera)
  • Nematoida (Nematoda and Nematophora)
  • Panarthropoda (Tardigrada, Onychophora and Euarthropoda)
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8
Q

Describe what is understood about Lobopodia (and name some examples)

A

“Worms with legs” - many seen in NOT a monophyly -> probably plesiomorphic ancestor state of MANY panarthropods - may resemble Panarthropod Common Ancestor

Many had an armoured and arthropod-like digestive system

Hallucigenia, Megadictyon, Diania, Xenusion, Jianshanopodia

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9
Q

Describe Opabinia and its significance

A

Opabinia is a possible stem-Arthropod from the Cambrian Period (probably between Megadictyon and Anomalocaridids on the tree)

Appears like a lobopodian but with swimming appendages and euarthropod-like compound eyes

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10
Q

Describe Megadictyon and its significance

A

A possible stem-Arthropod, probably more “stemward” than Opabinia and Anomalocaris

Appears like a lobopodian, considered a stem-Arthropod as it is armoured and has an Arthropod-like digestive system

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11
Q

Describe Anomalocaridids (e.g., Anomalocaris) and their significance

A

A possible Euarthropod stem group with arthropod-like jointed raptorial oral/feeding and swimming appendages

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12
Q

Describe the Bivalved Arthropods (e.g., Canadaspis)

A

Complex and specialised biramous appendages (like many euarthropods)

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13
Q

Describe the Megacheirans (Great Appendage Arthropods)

A

Have biramous swimming AND walking appendages along the whole body

Great appendages at front (crab/lobster like) - may be homologous with those of euarthropods - but hard to be sure

(e.g., genus Leanchoilia)

Some hypothesise that we can prove homology using fossilised brains -> debatable whether nerve tissue can be this well preserved?

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14
Q

Summarise the general trend(s) seen in Arthropod evolution

A

Along the stem lineage of Euarthropoda, we see a step-wise acquisition of arthropod-like traits:
- Compound eyes
- Arthropodised cephalic and trunk limbs
- Arthrodisation (appeared AFTER arthropodisation)
- Sternites and tergites

Probable order of key players:
Megadictyon -> Opabinia -> Anomalocaridids -> Bivalved Arhropods -> Great Appendage Arthropods -> Euarthropods

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15
Q

What is the main evidence for Lophotrochozoan monophyly?

A

Genetic Data - 18S ribosomal gene data, ESTs, 28S nuclear ribosomal gene, myosin II heavy chain

BUT no single synapomorphic morphological trait

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16
Q

Summarise our current understanding of Lophotrochozoan phylogeny

A

Lophotrochozoa are a reasonably well-established clade of around 13 phyla, named after the Lophophore (a feeding structure) and the Trochophore (larval morphology)

However, there is no common morphological trait across all their groups, and the internal resolution of this group’s phylogeny is controversial

Generally thought to be two major groups within Lophotrochozoa:
- one includes Mollusca, Annelida, Ribbon Worms (Nemertea), Phoronida and Brachiopoda
- one includes Bryozoa, Entoprocta, Cycliophora, the three Gnathiferan phyla (Gnathostomalida, Rotifera and Micrognathozoa), Gastrotricha and Platyhelminthes

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17
Q

Name and describe the “first” three Lophotrochozoan phyla within the non-Trochozoan group

A

BRYOZOA - ectoprocts, sessile, live in colonies, mainly marine

ENTOPROCTA - similar lifestyle to Bryozoa but different anus position [note: no proper lophophore]

CYCLIOPHORA - live on lobsters’ feeding parts (very limited niche)

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18
Q

Name the group of 3 Phyla grouped together in lecture, but limited evidence for monophyly in literature

A

Gnathifera (Gnathostomulida, Rotifera, Micrognathozoa)

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19
Q

Name and describe the 5 Phyla in the Platyzoa group

A

Gnathostomulida “jaw worms”

Rotifera - tiny, thorny-headed worms, some are parasitic

Micrognathozoans “jaw animals”

Gastrotricha “hairybacks” - worm-like

Platyhelminthes (includes flatworms, flukes and tapeworms)
-> most are parasitic
-> had a coelom but lost over time, so not as primitive as thought
-> eyespots and nervous tissue clusters at anterior, two nerve cords along body

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20
Q

Name and describe the three phylogenetically closest Lophotrochozoan species to the Annelids and Molluscs

A

Phoronida - Horseshoe worms - another sessile filter feeder

Brachiopoda - resemble bivalve molluscs (but this is purely convergent, different phyla!)

Nemertea - ribbon worms, can be very long, no segmentation, proboscis/stylet - often used in conjunction with neurotoxin

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21
Q

Describe the ecology of Annelida

A

Annelida (eg. earthworms and Hirundidae/leeches) can be marine or terrestrial, and can be sessile, predatory or take more bizarre forms

They are ecologically vital for fertilisation, aeration, drainage and preventing compaction

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22
Q

Describe the key feature of Annelida mentioned in the lecture

A

METAMERISM - segmentation in which morphological features are repeated in each segment - allows larger size while still moving efficiently

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23
Q

Describe the ecology of Mollusca

A

Most are herbivorous, though some (e.g., squids) are predatory

Includes Gastropods, Bivalves, Cephalopods and many more obscure ones

Second most speciose phylum in Animalia!

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24
Q

Describe the Gastropods

A
  • Includes Snails, Slugs, Sea Angels and Nudibranch
  • Often have an enlarged locomotive foot
  • Most (but not all) have a radula on top of the odontophore - buccal mass houses the protractor/retractor muscles
  • Most have a calcium carbonate shell
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25
Q

Describe the cephalopoda

A

Includes squid, octopus and others (e.g. pharaoh cuttlefish)

Many convergent traits with other aquatic animals, also with human eye

Octopuses highly intelligent - surprising given their ecology

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26
Q

What are the three main subphyla of arthropods?

A

Trilobites (extinct)
Chelicerates
Mandibulates

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27
Q

Describe the key features of arthropods

A
  • A segmented body with jointed appendages
  • Tagmatisation - segments fused together into functional regions (e.g. head, thorax, abdomen)
  • Highly cephalized (concentration of sensory organs at anterior end of body - often sophisticated mouthparts and sensory organs)
  • Open circulatory system
  • Chitinous exoskeleton that moults (ecdysis)
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28
Q

Describe the features that allowed Arthropods to be so successful

A
  • Segmented body allows efficient locomotion
  • Highly developed sense organs
  • Exoskeleton versatility
  • Complex learned AND unlearned behaviours
  • Tracheal system pumps air directly to cells
  • Metamorphosis allows exploitation of multiple niches by one species
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29
Q

Briefly describe the trilobites and their presence in the fossil record

A

The trilobites are an extinct subphylum of arthropods, with an exoskeleton divided into three lobes

They were extremely successful for hundreds of millions of years following the Cambrian Explosion, but disappeared relatively quickly in the Devonian

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30
Q

Describe the anatomy of chelicerates

A
  • They have segments (though these are not always obvious in adults)
  • Body is divided into two main regions (prosoma at front, opisthosoma at back)
  • Six appendage pairs on prosoma: chelicerae, pedipalps (unique to chelicerates!) and 4 walking limb pairs
  • The chelicerae can be scissor-like (e.g. Opiliones, Scorpiones, Xiphosura) OR highly specialised (e.g., fangs and venome in Aranae)
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31
Q

Discuss the reasons why the phylogeny of Chelicerates is difficult to resolve

A
  • Their origins are extremely ancient (500mya in middle Cambrian)
  • Rapid diversification of early Arachnids in Devonian
  • Extinction of some orders (e.g., Eurypterida/Sea Scorpions)
  • RAPID evolution in some Arachnid orders -> LBA
32
Q

Which Chelicerate Class is sister to all other chelicerates?

A

Pycnogonida

33
Q

Name the extant Chelicerate classes (and some key groups within them)

A
  1. Pycnogonida/Sea Spiders (Type Genus = Pycnogonum)
  2. Merostomata (includes extinct Eurypterida and extant Xiphosura - horseshoe crabs)
  3. Arachnida (includes Aranae, Scorpiones, Opiliones, Acari and others)
34
Q

What are the Eurypterida and what is the most striking feature about them compared to other arthropods?

A

They were sea scorpions, and an extinct group of Merostomata

Some were extremely large - possibly over 2m in size, making J. rhenaniae the largest known arthropod in history (ARTHROPOD GIGANTISM)

35
Q

What are the Xiphosura (brief anatomy and human significance)?

A

Horseshoe crabs - they have a wide abdomen and a telson (tailpiece); they are protected by a horseshoe-shaped, unsegmented dorsal shield (CARAPACE)

They are farmed for their blue blood (which contains Haemocynids rather than Haemoglobin, making it useful for clinical research)

36
Q

Name 7 of the 15 extant orders of Arachnids (plus common names) and 1 extinct class (and why it is significant)

A
  1. Araneae (spiders)
  2. Scorpionida (scorpions)
  3. Acari (mites + ticks)
  4. Opiliones (daddy-long-legs)
  5. Solifugae (sun/camel spiders)
  6. Thelyphonida (whip-scorpions)
  7. Amblypygi (tailless whip-scorpions)

Plus Trigonotarbida - possibly first ever animals to invade the land (computer graphics software used to simulate its movement; probably used an “alternating tetrapod gait”)

37
Q

Describe the segmentation seen in order Araneae

A

They have only TWO TAGMATA - the cephalothorax (a fused head and thorax) and the abdomen

Each of these shows no external segmentation, and they are joined by a narrow pedicel

38
Q

[Bit more on spiders if time]

A

Of course my guy! (salute emoji)

39
Q

Describe the anatomy and ecology of Scorpionida

A

They have a short cephalothorax with appendages, and a segmented abdomen with NO appendages (the sting is formed on the postabdomen and is usually harmless to humans)

Have many desert adaptations, e.g., slow metabolism and movement; the mother carries the (well-developed) young until their first moult

40
Q

Describe the Pycnogonida anatomy, ecology and phylogeny

A

Their head region (Cephalosoma) always bears a proboscis, and usually four eyes + four appendage pairs (the chelifores, palps, first leg pair and ovigers)
Organ systems are displaced in the legs (as the body is so slender)
Mostly marine; feed on sponges, cnidaria and even molluscs; some giant (unclear why) but most are small
Limited fossil record, but first appearance around 500mya in Middle Cambrian; generally agreed to be sister to all other chelicerates

41
Q

Describe the notable features of Opiliones

A

Daddy-long-legs have a single fused body segment and long legs
They show diverse feeding behaviours, including mastocation (chewing)
They are useful for biogeography due to their lack of cross-ocean migration

42
Q

Describe the notable features of Solifugae

A

SOLifugae (SUN spiders!) are generally quite large and predate on small arthropods
They are an indicator species of desert biomes

43
Q

[Maybe more on Thelyphonida and Amblypygi if time]

A

No worries fella (salute emoji)

44
Q

Which group best demonstrates prehistoric Arthropod Gigantism (and what are the possible reasons for this)?

A

Eurypterida (e.g., J. rhenaniae) - possible due to increased oxygen in the atmosphere, or due to an evolutionary arms race with prey (both debatable)

45
Q

Describe the notable features of Acari

A

Mites + Ticks - monophyly is possible but debated

They are very diverse and found in virtually all habitats; many are parasitic and have major human health implications (e.g., Lyme Disease, Babesiosis, Scabies)

46
Q

Summarise the phylogenetic relationships between the major clades of arthropods (and how this has dramatically changed in recent years)

A

Chelicerates are sister to other arthropods - the other major groups are collectively called “Mandibulata”

Mandibulata includes Myriapoda and Pancrustacea

Pancrustacea INCLUDES insecta, showing that “crustaceans” are paraphyletic

47
Q

Describe the groups within the Myriapoda

A

Four extant classes:
- Chilopoda (e.g. centipedes) is sister taxon
- Symphyla (not very speciose, mainly detritus-feeding soil inhabitants)
- Pauropoda (see above)
- Diplopoda (e.g. millipedes) is most speciose

48
Q

Describe the notable features of Diplopoda

A

Diplopoda = Millipedes

They have 2 pairs legs per segment (with those of segment 7 being modified into reproductive organs, Gonopods)
Also have a CaCO3 exoskeleton
They feed on dead plant material and may have been the first land animals (415mya feces)
Includes possibly the largest land arthropod ever (Arthropleura)

49
Q

Describe the notable features of Chilopoda

A

Chilopoda (Centipedes) have just 1 walking pair per segment, and chelicerae modified into fangs (FORCIPULES) a bit like spiders

Very diverse ecology (e.g., large bat-eating cave centipedes), including maternal care!

50
Q

Name the Cambrian species believed to represent the first Pancrustacean fossil

A

Waptia fieldensis

51
Q

Name the 5 main (described) classes of Crustaceans from least- to most-closely-related to insects

A

Ostracoda - Thecostraca - Malacostraca - Branchiopoda - Remipedia

52
Q

Describe the notable features of Ostracoda

A

Resemble Camrbian bivalve arthropods (though unclear if related)

Diverse ecology, tolerant of many conditions, radiated in Silurian into many forms of plankton

53
Q

Describe the notable features of Thecostraca

A

Adult forms all sessile (parasites OR suspension feeders like Cirripeda, which includes barnacles)

54
Q

Describe the notable features of Malacostraca

A

Includes lobsters, crabs, isopods (many archetypal crustaceans)

20 body segments

55
Q

Describe the notable features of Branchiopoda

A

Include Daphnia (recognisable zooplankton), inducible defenses

56
Q

Describe the notable features of Remipedia

A

Blind, cave-dwelling, tropical, no carapace (hard covering), closely related to Insecta

57
Q

Describe the life cycle of a typical crustacean

A
  1. Egg
  2. Nauplius (hard to identify, few pass this stage)
  3. Protozoea
  4. Mysis
  5. Postlarval
  6. Adult
58
Q

Describe the order Isopoda

A

One of around 15 orders within class Malacostraca; 5000 species, mainly marine; Abyssal Gigantism (large in deep sea due to pressure)

No carapace, dorsally flattened, lack effective cuticle

Females carry young in MANCAE until hatch

59
Q

Describe the order Decapoda

A

One of around 15 orders within class Malacostraca; includes crabs and lobsters!

2 pairs of antennae, 5 pairs of walking legs

Radiated in Triassic (CRABS radiated in Jurassic) - note: carcinisation

60
Q

Name the three orders of non-Insect Hexapods

A

Protura, Diplura and Collembola (their exact phylogney is unclear)

61
Q

Describe the general anatomy of the class Insecta

A

Body divided into three segments:

  • Head
  • Thorax (which is further divided into segments T1, T2 and T3)
  • Abdomen

[Fore- and hind-wings generally found on T2 and T3, while one pair of walking legs is found on each Thoracic segment; Abdomen is generally segmentless]

Spiracles for ventilation, compound eye, reproductive organs on abdomen, Cercus at back of abdomen - adapted for various functions, including clasping in reproduction

62
Q

Name and briefly describe the main synapomorphy of Insecta

A

Johnston’s organ - a sensory organ involved in sound detection, located in the second antennal segment

63
Q

Why are insects thought to be so successful and resistant to extinction? (Both general trends and specific traits)

A

They have high speciation rates and reproductive capacity, short generation times, and significant genetic diversity, all of which allows them to rapidly adapt to new environments or pressures

Also an amalgamation of traits that work synergistically, e.g.:

  • Labile segmented body plan
  • Resilient arthropod exoskeleton
  • Flight and neopterous wings (able to escape predators and move to new environments)
  • Herbivory
  • Larval stage and holometaboly (exploit multiple niches)
64
Q

Describe what is meant by “hemimetabolous” and “holometabolous”

A

Hemimetabolous: No pupal stage in development - generally gradual growth, with each instar resembling the last [Exopterygota]

Holometabolous: Larval and Pupal stages; often extreme transformation, allowing them to exploit multiple niches [Endopterygota]

65
Q

Given that insects are the most successful group on land, and some are even aquatic, why are there no marine insects?

A

No clear answer - possible factor is that crustaceans had already colonised every niche and insects could not outcompete them

Maddrell (1998) argued that fish predation is the main factor - insects cannot escape fish predation by vertical migration, as many crustaceans do, because their tracheal system of ventilation only functions in the top 30-40m of water

66
Q

When were the first (potential) insects, and how likely are these fossil to be genuine insects?

A

Rhyniognatha hirstii - single early Devonian (400mya) fossil [Haug and Haug (2017) reinterpreted some key features of this fossil, including the mandibles, and consequently argued that the fossil could be a centipede or other myriapod]

Rhyniella praecursor - Lower Devonian, more complete fossil, generally accepted, contaminant theory was dismissed in 1988

Strudiella devonica - discovered in 2012 and interpreted as an insect, shrinking the Hexapod gap in the fossil record. However, now its insect status is disputed and it can only be agreed to be an Arthropod of some form

67
Q

What is the Hexapod gap?

A

A long period in the fossil record with very few Hexapod fossils, lasting from the late Devonian to the early Carboniferous

This is largely due to the poor preservation of insects due to their size and anatomy

(The gap may have been narrowed significantly in 2012 by the discovery of Strudiella devonica, but its status as an insect was disputed by Hörnschemeyer et al in 2013)

68
Q

What is significant about the insect order Palaeodictyopterida?

A

It is the only known insect order to have gone extinct (died out in the end-Permian, around 250mya, unclear why)

Palaeodictyopterida were beaked, medium-to-giant insects (linked to increased O2 levels), related to Odonata, possibly the first major group of herbivorous insects

69
Q

What do the Xylocopinae reveal about insect evolution (and try to name the four tribes for bonus points)?

A

The four tribes of long-tongued bees (Xylocopinae) are Manuellini, Xylocopini, Ceratinini and Allodapini

When their phylogeny is displayed as a radiating diagram (see slides), there are long black lines, before relatively sudden radiation of the four tribes

This could suggest “long fuse” - very little speciation for a long time OR it could suggest mass extinctions

70
Q

What are the two main subclasses of insects?

A

Pterygota (winged insects) and Apterygota (wingless insects)

NOTE: Apterygota is a disused and paraphyletic taxon

71
Q

Describe the Colyoptera

A

BEETLES - most diverse order in animal kingdom!

First appeared 265mya, elytron (hard outer wing shell), 95% are herbivorous - possible causality?

McKenna et al (2019) suggest that success of beetles is not linked to herbivory itself, but rather to genomic innovation and endogenous Plant Cell Wall Degrading Enzymes (PCWDEs)

72
Q

Give the common name for the Order Mecoptera

A

Scorpionflies (not scorpions or flies!)

73
Q

Describe the order Odonata

A

DRAGONFLIES AND DAMSELFLIES - spend most of their life cycle underwater, then final stage in the air

Emerged in Carboniferous; carnivorous

Dragonflies hold wings perpendicular, damselflies hold wings parallel

74
Q

Describe the order Hymenoptera

A

BEES, WASPS AND ANTS - Have a synapomorphy: hamuli (wing claspers)

Emerged around 281 MYA (Permian?)

Eusociality and also many key pollinators

Ants (Formicidae) in particular are found on every continent and have a huge biomass

75
Q

Describe the order Dyptera

A

FLIES - important in human disease (e.g., mosquitos), genetics (e.g., Drosophila) and pollination

First appeared in Triassic (crane fly)

76
Q

Describe the notable features of the subclass Cirripeda

A

Filter-feeding Crustacean sub-class within class Thecostraca (includes the Barnacles, which have the longest penises relative to body size in the animal kingdom)

77
Q
A