Alexander Wild (all)

Virtual Entomology for Master Naturalists Rivanna Master Naturalists 8 April 2020

Linda S. Fink Duberg Professor of Ecology Sweet Briar College Many of today’s beautiful images were taken by Alex Wild (https:// www.alexanderwild.com/)

Goals for Insect Day Understand 1. features that characterize arthropods in general, insects specifically 2. how much variation there is in all aspects of insect biology 3. ecological importance of insects

Feel prepared to 4. participate in insect projects as Master Naturalists • children's and school programs • pollinator and foodplant gardening • monitor native insect populations, stream health, exotic insects • measure biodiversity (e.g. bioblitz, NABA butterfly count) • citizen science (e.g. Journey North, Monarch Watch) 5. learn more about insects

Structure of the presentation

PowerPoint 1. Introduction to arthropods and insects

break

PowerPoint 2. Why are insects so successful? break

PowerPoint 3. The seven largest groups of insects Insects are the dominant multicellular life form on the planet

A “species scape” • Number of species • Numbers of individuals • Biomass

Why learn about insects?

"the insects are so numerous that if they were divided equally among each one of the earth's 6 billion human inhabitants, each of us would be allotted 1 x 1018 insects -- that's a billion billion -- 1,000,000,000,000,000,000." J. Myers, some years ago

mayflies

locusts monarch butterflies Why learn about insects?

Human requirements Agriculture and food production beneficial and harmful Health and disease human, livestock, companion animals, plants Scientific discovery Culture Economics

Why learn about insects?

Ecological roles pollination phytophagy seed dispersal fungal dispersal nutrient cycling predators and parasites prey Bumblebee buzz pollinating a tomato blossom

Peponapis squash bee Andrenid bee on an apple blossom Why learn about insects?

Ecological roles pollination phytophagy seed dispersal fungal dispersal stem borer nutrient cycling predators and parasites phyto- plant prey fruit pest phag(o)- eat

leaf miner phloem feeder Why learn about insects?

Ecological roles pollination phytophagy seed dispersal fungal dispersal nutrient cycling predators and parasites prey

But the best reason to study insects is...

But the best reason to study insects is...

they are amazing

Alex Wild But the best reason to study insects is...

they are amazing

Alex Wild But the best reason to study insects is...

they are amazing

Alex Wild But the best reason to study insects is...

they are amazing

Alex Wild 2018 headlines Hallman, C.A. et al. 2017. More than 75 percent decline over 27 years in total flying insect biomass in protected areas. PLoS ONE 12(10): e0185809.

Sanchez-Bayo, F. and K.A.G. Wyckhuys. 2019. Worldwide decline of the entomofauna: A review of its drivers. Biol Cons 232: 8-27. F. Sánchez-Bayo, K.A.G. Wyckhuys Proportion of terrestrial insect species in decline or locally extinct A) Terrestrial taxa

decline <30% vulnerable endangered exnct

1 0.9 0.8 0.7 0.6 0.5 0.4 0.3 Propor  on of species 0.2 0.1 0

B) Aquac taxa 0.8 Sanchez-Bayo & Wyckhuys. 2019. Biological Conservation 232: 8-27. 0.7

0.6

0.5 exnct

0.4 endangered vulnerable 0.3 decline <30% Propor  on of species 0.2

0.1

0.0 Ephemeroptera Odonata Plecoptera Trichoptera

Fig. 3. Proportion of insect species in decline or locally extinct according to the IUCN criteria: vulnerable species (> 30% decline), endangered species (> 50% decline) and extinct (not recorded for > 50 years). A) terrestrial taxa; B) aquatic taxa.

Davis et al., 2004 ; Kreutzweiser et al., 2007 ). services, but it's unclear to what extent natural ecosystems can sustain While countless insect species are disappearing, few others are oc- their overall ecological resilience (Memmott et al., 2004 ). cupying vacant niches and expanding their distribution. In terrestrial Species extinctions equally impact the overall biomass of entire ecosystems, most of the occupying species are generalists with diverse ecosystems, as insects form the base that supports intricate food webs. ecological preferences (e.g., Bombus impatients, Plusia putnami, Indeed, the essential role that insects play as food items of many ver- Laemostenus terricola and Hippodamia variegata). In aquatic environ- tebrates is often forgotten. Shrews, moles, hedgehogs, anteaters, lizards, ments, species replacement is also mediated by ecological traits such as amphibians, most bats, many birds and fish feed on insects or depend degree of tolerance to pollutants (e.g. Sympetrum striolatum, Brachyptera on them for rearing their offspring. Even if some declining insects might risi and Potamyia flava), with communities thus becoming more uniform be replaced with others, it is difficult to envision how a net drop in and less diverse in composition (Houghton and Holzenthal, 2010). overall insect biomass could be countered. The large declines in insect Species replacement may help retain the delivery of certain ecosystem biomass observed in Europe (Hallmann et al., 2017 ) and Puerto Rico

Proportion of declining insect species F. Sánchez-Bayo, K.A.G. Wyckhuys

Box Mean line Mild outliers (agriculture) and manufacture goods (industrialisation) at the expense 1 n = 34 n = 11 n = 15 n = 9 of various natural habitats. Among Coleoptera, Lepidoptera and 0.9 Hymenoptera, land-use change and landscape fragmentation is surely the main cause of species declines (Fig. 5), with agricultural conversion 0.8 and intensification for food production listed in 24% of the reports 0.7 (Fig. 6). Urbanisation, by contrast, is reported in 11% of cases, while 0.6 deforestation appears in 9% of reports. 0.5 As agricultural crops comprise about 12% of the total land surface 0.4 on the planet (FAO, 2015), farming directly affects a considerable 0.3 proportion of insect species (Dudley and Alexander, 2017). In Europe 0.2 and North America, the expansion of the agricultural frontier took place fi

on of declining species declining of  on Propor mostly in the rst half of the 20th century, whereas in South America, 0.1 Africa and Asia occurred mainly in the second half of the century (Foley 0 et al., 2005; Gibbs et al., 2010). In its wake, rare species associated with Europe U.K. N America other pristine ecosystems and natural habitats either retreated or were en- Fig. 4. Proportion of declining insect species in different regions of the world. tirely lost (Grixti et al., 2009; Ollerton et al., 2014). Major insect de- clines occurred, however, when agricultural practices shifted from traditional, low-input farming style to the intensive, industrial scale (Lister and Garcia, 2018) inevitably lead to a starvation of dependent Sanchez-Bayo & Wyckhuys. 2019. production brought about by the Green Revolution (Bambaradeniya vertebrates (Hallmann et al., 2014; Lister andBiological Garcia, Conservation 2018; Poulin 232: 8-27. and Amerasinghe, 2003; Ollerton et al., 2014). The latter practices did et al., 2010; Wickramasinghe et al., 2003). This kind of cascading effect not necessarily involve deforestation or habitat modification (e.g., was first observed with grey partridge (Perdix perdix) populations in grassland conversion, drainage of wetlands) but rather entailed the England since 1952, and was ascribed to reproductive failure. The ul- planting of genetically-uniform monocultures, the recurrent use of timate cause of the partridge collapse was a combined use of in- synthetic fertilisers and pesticides, the removal of hedgerows and trees secticides and herbicides in agricultural land, leading to insufficient in order to facilitate mechanization, and the modification of surface insect numbers to feed the chicks (Potts, 1986). Equally, in the U.K. the waterways to improve irrigation and drainage. Monocultures led to a diversity and abundance of bats in intensive agricultural landscapes is great simplification of insect biodiversity among pollinators, insect considerably lower than on organic farms because of a reduction in natural enemies and nutrient recyclers, and created the suitable con- insect biomass caused by pesticide use in the former settings ditions for agricultural pests to flourish. A quarter of the reports in- (Wickramasinghe et al., 2004), and direct insecticide exposure through dicate these agriculture-related practices as the main driver of insect the bats' prey items (Mispagel et al., 2004; Stahlschmidt and Bruhl, declines in both terrestrial and aquatic ecosystems (Wilcove et al., 2012). 1998). The susceptibility of specialist pollinators to land-use changes (in- 4.1. Drivers of the declines volving loss of floral resources, nesting and hibernation sites), appears to be a determining factor in the decline of many bumblebees and wild A large proportion of studies (49.7%) point to habitat change as the bees (Williams and Osborne, 2009). For specialist ground beetles, the main driver of insect declines, a factor equally implicated in global bird loss of hedgerows and trees likely triggered their decline (Brooks et al., and mammal declines (Chamberlain and Fuller, 2000; Diamond, 1989). 2012). Declines in moths are tied to the fate of their overwintering Next on the list is pollution (25.8%) followed by a variety of biological larval host plants: forbs for species overwintering as larvae, and trees factors (17.6%), whereas few studies (6.9%) indicate climate change as for those overwintering as egg, pupa, or adult. The combined removal triggering the losses (Fig. 5; Table S2). of weeds and trees in intensive agricultural settings may thus explain the decline of moth species overwintering as larvae (Fox, 2013; Mattila 4.1.1. Habitat change et al., 2006; Merckx et al., 2009; Pocock and Jennings, 2008). Con- Habitat change is an immediate consequence of human activities. Its versely, the change from intensive farming to organic farming has led to global pace and scope has been expanding over the past centuries, with increases in abundance and diversity of moths (Taylor and Morecroft, increasing amounts of land being transformed to provide dwellings, 2009), while the abandonment of grazing land has allowed the recovery facilitate transportation and enable tourism (urbanisation), grow food of some common butterflies (Kuussaari et al., 2007). Agricultural intensification also entails stream channelization, 90 draining of wetlands, modification of floodplains, and removal of ri- 80 parian canopy cover with subsequent loss of soil and nutrients – all resulting in homogenization of stream microhabitats and alteration of 70 aquatic insect communities (Houghton and Holzenthal, 2010). These 60 Coleoptera activities increase eutrophication, siltation and sedimentation in water Hymenoptera bodies, thus reducing the richness of shredders and predators while 50 fi Lepidoptera favouring lterer species (Burdon et al., 2013; Niyogi et al., 2007; Olson 40 Odonata et al., 2016). Diverse communities of aquatic plants are an important habitat component in lentic systems such as paddy fields, allowing

Number of reports Other aquac 30 herbivory, oviposition and emergence of many insects and providing Other terrestrial 20 refugia for Odonata nymphs (Nakanishi et al., 2014). In general, loss of permanent flows in streams and rivers leads to a decrease of biodi- 10 versity (King et al., 2016), whereas irrigation and man-made water 0 bodies in urbanised areas may have favoured certain species (Kalkman Habitat change Polluon Biological traits Climate change et al., 2010). In recent decades, urbanisation has taken over agricultural land Fig. 5. The four major drivers of decline for each of the studied taxa according across the globe, causing the disappearance of many habitat specialists to reports in the literature. and their replacement with a few generalists adapted to the artificial

F. Sánchez-Bayo, K.A.G. Wyckhuys Major causes of insect declines

1.3% 1.9% 1.9% 1.9% 3.1% intensive agriculture pescides

5.0% 23.9% ecological traits urbanisaon 6.3% ferlisers deforestaon

8.8% wetlands/rivers alteraon warming 12.6% other pollutants

10.1% pathogens fires 12.6% introduced species 10.7% genec

Fig. 6. Main factors associated with insect declinesSanchez-– see alsoBayoFig. & 5 .Wyckhuys. 2019. Biological Conservation 232: 8-27. human environment. However, such losses can be partially counter- Herbicides, however, reduce the biodiversity of vegetation within the balanced by the creation of urban parklands and gardens, which offer crops and in surrounding areas through drift (Egan et al., 2014) and refuge to native and newly-colonising species, including pollinators like runoff, thus impacting indirectly on the arthropod species that depend Bombus spp. (Botías et al., 2017) and butterflies like Lycaena phlaeas upon wild plants, which either disappear completely or decline sig- and Aphantopus hyperantus (van Dyck et al., 2009). nificantly in numbers (Goulet and Masner, 2017; Marshall et al., 2003). In tropical countries of South America, Africa and Asia, deforesta- Thus, the application of herbicides to cropland has had more negative tion has been and still is a main driver of biodiversity loss and insect impacts on both terrestrial and aquatic plants and insect biodiversity declines (Carrasco et al., 2017; Wilson, 2002), including dragonflies than any other agronomic practice (Hyvonen and Salonen, 2002; (Samways, 1999). Recent research on herbariums of Pacific islands Lundgren et al., 2013). Pesticides have caused the decline of moths in suggests that deforestation and other human impacts on those ecosys- rural areas of the U.K. (Hahn et al., 2015; Wickramasinghe et al., 2004) tems are not confined to the extinction of birds, mammals and snails and pollinators in Italy (Brittain et al., 2010); broad-spectrum in- (Kingsford et al., 2009) but also of insects such as leafminers (Lepi- secticides reduce the abundance and diversity of beneficial ground- doptera: Gracillariidae) (Hembry, 2013). In Europe, deforestation is the dwelling and foliage-foraging insects (Lundgren et al., 2013); systemic main driver of saproxylic beetles' decline (Nieto and Alexander, 2010). insecticides reduce populations of ladybirds and butterflies in gardens Conversely, afforestation may increase the number of generalist but- and nurseries (Krischik et al., 2015), and inflict multiple lethal and sub- terflies by increasing habitat diversity at the forest edge (Kuussaari lethal effects on bees (see 3.2.3) and other arthropods. Fungicides are et al., 2007), but woodland diversity, structural and micro-climatic not less damaging to insects, and synergism of a particular group of heterogeneity are far more important than forested area per se for compounds (i.e., azoles) with insecticide toxicity (Biddinger et al., maintaining the diversity of moths, butterflies as well as birds (Fuller 2013; Pilling and Jepson, 1993) is certainly involved in honey bee et al., 2005; van Swaay et al., 2006). Very few generalist species benefit collapses (Simon-Delso et al., 2014). and expand under afforestation, and some European butterfly species Pyrethroid, neonicotinoid and fipronil insecticides have a devas- even exhibited notable declines (van Swaay et al., 2006). In Britain, a tating impact on aquatic insects and crustaceans due to their high acute 20-fold increase in conifer plantations since the 19th century did not and chronic toxicity (Beketov and Liess, 2008; Kasai et al., 2016; Mian increase biodiversity nor abundance of Lepidoptera species (Brereton and Mulla, 1992; Roessink et al., 2013), thus reducing significantly et al., 2011; Fox, 2013). their abundance in water bodies (van Dijk et al., 2013). Persistent re- sidues of fipronil in sediments inhibit the emergence of dragonflies (Jinguji et al., 2013; Ueda and Jinguji, 2013) and the development of 4.1.2. Pollution chironomids and other insect larvae, with negative cascading effects on Pollution is the second major driver of insect declines (Fig. 5). fish survival (Weston et al., 2015). Systemic insecticides impair the Sources of environmental pollution include fertilisers and synthetic long-term viability of shredder larvae that decompose leaf litter and pesticides used in agricultural production, sewage and landfill leachates other organic material (Kreutzweiser et al., 2008), undermine the basis from urbanised areas and industrial chemicals from factories and of the insect food web (Sánchez-Bayo et al., 2016a) and thus derail mining sites. Among these, pesticide pollution is reported in 13% of natural biological control mechanisms e.g., in rice paddy ecosystems cases (Fig. 6), followed by fertiliser inputs (10%) and to a lesser extent (Settle et al., 1996). Also, these products readily translocate to pollen, urban and industrial pollutants (3%). nectar, guttation drops, and all tissues of the treated crops and adjacent Intensive agriculture implies the systematic and widespread use of plants, impacting on nectar-feeding biota such as bees, butterflies, ho- pesticides for controlling crop pests (insecticides), competing weeds verflies and parasitic wasps (van der Sluijs et al., 2015). Unlike the (herbicides) and fungal infections (fungicides) among others (Dudley short-term effects of other pesticides on aquatic organisms (Schäfer and Alexander, 2017). In terms of toxicity, insecticides are by far the et al., 2011; van den Brink et al., 1996), neonicotinoids do not allow the most toxic to all insects and other arthropods, followed by fungicides recovery of univoltine and semivoltine aquatic insects (Beketov et al., but not herbicides (Mulé et al., 2017; Sánchez-Bayo and Goka, 2014).

F. Sánchez-Bayo, K.A.G. Wyckhuys Major causes of insect declines

1.3% 1.9% 1.9% 1.9% 3.1% intensive agriculture pescides

5.0% 23.9% ecological traits 36%urbanisa on 6.3% Intensiveferlisers agriculturedeforestaon 8.8% andwetlands/rivers Pesticides alteraon warming 12.6% other pollutants

10.1% pathogens fires 12.6% introduced species 10.7% genec

Fig. 6. Main factors associated with insect declinesSanchez-– see alsoBayoFig. & 5 .Wyckhuys. 2019. Biological Conservation 232: 8-27. human environment. However, such losses can be partially counter- Herbicides, however, reduce the biodiversity of vegetation within the balanced by the creation of urban parklands and gardens, which offer crops and in surrounding areas through drift (Egan et al., 2014) and refuge to native and newly-colonising species, including pollinators like runoff, thus impacting indirectly on the arthropod species that depend Bombus spp. (Botías et al., 2017) and butterflies like Lycaena phlaeas upon wild plants, which either disappear completely or decline sig- and Aphantopus hyperantus (van Dyck et al., 2009). nificantly in numbers (Goulet and Masner, 2017; Marshall et al., 2003). In tropical countries of South America, Africa and Asia, deforesta- Thus, the application of herbicides to cropland has had more negative tion has been and still is a main driver of biodiversity loss and insect impacts on both terrestrial and aquatic plants and insect biodiversity declines (Carrasco et al., 2017; Wilson, 2002), including dragonflies than any other agronomic practice (Hyvonen and Salonen, 2002; (Samways, 1999). Recent research on herbariums of Pacific islands Lundgren et al., 2013). Pesticides have caused the decline of moths in suggests that deforestation and other human impacts on those ecosys- rural areas of the U.K. (Hahn et al., 2015; Wickramasinghe et al., 2004) tems are not confined to the extinction of birds, mammals and snails and pollinators in Italy (Brittain et al., 2010); broad-spectrum in- (Kingsford et al., 2009) but also of insects such as leafminers (Lepi- secticides reduce the abundance and diversity of beneficial ground- doptera: Gracillariidae) (Hembry, 2013). In Europe, deforestation is the dwelling and foliage-foraging insects (Lundgren et al., 2013); systemic main driver of saproxylic beetles' decline (Nieto and Alexander, 2010). insecticides reduce populations of ladybirds and butterflies in gardens Conversely, afforestation may increase the number of generalist but- and nurseries (Krischik et al., 2015), and inflict multiple lethal and sub- terflies by increasing habitat diversity at the forest edge (Kuussaari lethal effects on bees (see 3.2.3) and other arthropods. Fungicides are et al., 2007), but woodland diversity, structural and micro-climatic not less damaging to insects, and synergism of a particular group of heterogeneity are far more important than forested area per se for compounds (i.e., azoles) with insecticide toxicity (Biddinger et al., maintaining the diversity of moths, butterflies as well as birds (Fuller 2013; Pilling and Jepson, 1993) is certainly involved in honey bee et al., 2005; van Swaay et al., 2006). Very few generalist species benefit collapses (Simon-Delso et al., 2014). and expand under afforestation, and some European butterfly species Pyrethroid, neonicotinoid and fipronil insecticides have a devas- even exhibited notable declines (van Swaay et al., 2006). In Britain, a tating impact on aquatic insects and crustaceans due to their high acute 20-fold increase in conifer plantations since the 19th century did not and chronic toxicity (Beketov and Liess, 2008; Kasai et al., 2016; Mian increase biodiversity nor abundance of Lepidoptera species (Brereton and Mulla, 1992; Roessink et al., 2013), thus reducing significantly et al., 2011; Fox, 2013). their abundance in water bodies (van Dijk et al., 2013). Persistent re- sidues of fipronil in sediments inhibit the emergence of dragonflies (Jinguji et al., 2013; Ueda and Jinguji, 2013) and the development of 4.1.2. Pollution chironomids and other insect larvae, with negative cascading effects on Pollution is the second major driver of insect declines (Fig. 5). fish survival (Weston et al., 2015). Systemic insecticides impair the Sources of environmental pollution include fertilisers and synthetic long-term viability of shredder larvae that decompose leaf litter and pesticides used in agricultural production, sewage and landfill leachates other organic material (Kreutzweiser et al., 2008), undermine the basis from urbanised areas and industrial chemicals from factories and of the insect food web (Sánchez-Bayo et al., 2016a) and thus derail mining sites. Among these, pesticide pollution is reported in 13% of natural biological control mechanisms e.g., in rice paddy ecosystems cases (Fig. 6), followed by fertiliser inputs (10%) and to a lesser extent (Settle et al., 1996). Also, these products readily translocate to pollen, urban and industrial pollutants (3%). nectar, guttation drops, and all tissues of the treated crops and adjacent Intensive agriculture implies the systematic and widespread use of plants, impacting on nectar-feeding biota such as bees, butterflies, ho- pesticides for controlling crop pests (insecticides), competing weeds verflies and parasitic wasps (van der Sluijs et al., 2015). Unlike the (herbicides) and fungal infections (fungicides) among others (Dudley short-term effects of other pesticides on aquatic organisms (Schäfer and Alexander, 2017). In terms of toxicity, insecticides are by far the et al., 2011; van den Brink et al., 1996), neonicotinoids do not allow the most toxic to all insects and other arthropods, followed by fungicides recovery of univoltine and semivoltine aquatic insects (Beketov et al., but not herbicides (Mulé et al., 2017; Sánchez-Bayo and Goka, 2014).

“If all mankind were to disappear, the world would regenerate back to the rich state of equilibrium that existed ten thousand years ago. If insects were to vanish, the environment would collapse into chaos.”

E. O. Wilson

Goals for Insect Day Understand 1. features that characterize arthropods in general, insects specifically 2. how much variation there is in all aspects of insect biology 3. ecological importance of insects

Feel prepared to 4. participate in insect projects as Master Naturalists • children's and school programs • pollinator and foodplant gardening • monitor native insect populations, stream health, exotic insects • measure biodiversity (e.g. bioblitz, NABA butterfly count) • citizen science (e.g. Journey North, Monarch Watch) 5. learn more about insects

Traditional Classification

Kingdom ANIMAL Phylum ARTHROPOD Class INSECT Order Family Genus Species Traditional Classification Kingdom: Animal Phylum: Major body plans

sponges chordates segmented worms molluscs roundworms echinoderms Let's look at a few body plans Let's look at a few These are ARTHROPODS body plans Phylum: Arthropods

centipedes spiders insects scorpions

crustaceans

horseshoe crab and more... Arthropod characteristics

•! Jointed, paired appendages •! External hardened skeleton of cuticle •! Periodically shed exoskeleton to grow (molt, ecdysis) •! Segmented body is organized into 2 or 3 functional units •! More than two pairs of limbs

Let’s look at the external skeleton Functions Physical protection Waterproofing Sites for muscle attachment Locations for sensory receptors Variation in exoskeletons

all photos from Alex Wild Variation in exoskeletons Honeypot ants (Camponotus inflatus)

dorsal sclerite membrane

Alex Wild Sensory structure examples: Chemoreceptors Molting (ecdysis)

•!External skeleton is not cellular •!it doesn’t grow •!To increase in size, an arthropod must •!shed its old skeleton •!produce a larger one

britannica.com

nutmeg66 on flickr.com

pinebaskets.tripod.com Molting (ecdysis) is risky: physical injury desiccation predation

www.arkinspace.com stuant63 Molting is associated with both growth and development Molting is associated with both growth and development

MOLTS Molting is associated with both growth and development

GROWTH Insects do most of their growth in the last immature instar

Hornworm caterpillars Manduca

early 5th instar late 5th instar Molting is associated with both growth and development

DEVELOPMENT Changes in form are associated with molting

Becky Hansis O’Neill Phylum: Arthropods

centipedes spiders insects scorpions

crustaceans

horseshoe crab and more... Arthropod characteristics

•! Jointed, paired appendages •! External hardened skeleton of cuticle •! Periodically shed exoskeleton to grow (molt, ecdysis) •! Segmented body is organized into 2 or 3 functional units •! More than two pairs of limbs

Traditional Classification

Kingdom ANIMAL Phylum ARTHROPOD Class INSECTA Order Family Genus Species What are the classes of Arthropods? Kingdom: Animal Phylum: Arthropod Class Insects • Three body regions: • head, thorax, abdomen • One pair of antennae • Three pairs of legs

photo by Alex Wild Among arthropods, flight occurs only in adult insects

Nicholl Williams

Stephen Dalton

photomacrography.net Non-insect Arthropods: Crustaceans •!two body regions: cephalothorax and abdomen •!two pairs of antennae •!most appendages are two-branched Non-insect Arthropods: Chelicerates

•!Arachnids (spiders, mites, ticks, scorpions)

•! plus a few other odd groups Non-insect Arthropods: Chelicerates • two body regions: jumping spider • cephalothorax and abdomen • lack antennae

• six pairs of unbranched pedipalp appendages • 1st appendage pair = chelicerae (jaws) chelicera • 2nd appendage pair = pseudoscorpion pedipalps These may appear leglike or may be highly modified • pairs 3-6 = leglike

pedipalp Non-insect Arthropods: Myriapods • Two body regions: head and trunk • One pair of antennae • Appendages unbranched • Many pairs of legs

• Millipedes: most are herbivores or detritivores • Centipedes: most are carnivores Let’s take a break